The Thirdof Agriculture,
Forestry(MAFF)

Ministry

International

and Fisheries,

Japan

Workshop on Genetic Resources

Animal Genetic

Resources:

Efficientand Effective

Conservation

Agriculture,

Forestry

Tsukuba Officeand Fisheries Research Council

Use

Secretariat

December 5 - 7, 1995

Sponsored by
Research Council Secretariat
andNational Instituteof Agrobiologicalof MAFF
Resources

in cooperation with
National Instituteof Animal Industry,
National Instituteof Sericultural& Entomological
Sciences,
National Instituteof Animal Health,andJapanese International
Research Center for Agricultural
SciencesContents
Page
Introduc i on
Welcome address
Opening

T. Hosoda

3

M.Nakagahra

5

address

Keynote addresses
Why and Howto Conserve DomesticatedAnimal Genetic Resources

History and Phytogeny of Japanese Native Animals and Strategies
Their Effective Use

I. Bodo7

Y.Tanabe

17
35

Ts.Zhanchiv

39

for

Discussion
Session 1 : Economic
Economic Evaluation
Economic Evaluation

evaluationof Mongolian Sheep Genetic Resourcesof Pig Genetic Resources in China

A.WangCharacterization
and Conservation of Silkworm Genetic Resources in India
RK.Datta
Chicken Genetic Resources in Japan and New Evaluation Research
K. Nirasawa, H.Takahashi, T. Furukawa, K. Kikuchi,
J. Noguchi, Y. Izaike and T. Oishi
Discussion
Session2:
Genetic
Genetic Classification
Polymorphisms

classificationof Asian Buffaloes

Based on Biochemical

Relationships

and Conservation

55

73
83

and Molecular
S.G.TanPhylogenetic
in Asia45

of Bovine and Bubaline

87

SpeciesT. Namikawa, K. Tanaka and C. D. Solis

99

The Phylogenetic
Relationship
Pigs (Sus scrofa L.) in Asia

and Differentiationof Wild and DomesticatedY.Kurosawa

111
12 1

L.V.Ly125

Discussion
Session 3: Conservation
systems
Animal Genetic Resources and Conservation
Conservation
Collection

Evaluation
Discussion

System in Vietnam

Systems of Animal Genetic

Resources in The Philippines
J.M.Matias
131
and Preservation of Animal Genetic Resources in the MAFF Gene Bank
Y. Izaike, K. Kikuchi, J. Noguchi, K. Nirasawa,
H. Takahashi, T. Furukawa and T. Oishi
141
and Preservation of Silkworm Genetic Resources in Japan
E. Kosegawa, T. Kikawada, K. Shimizu and T. Okajima
153
1 63

Session 4: New technology for conservation
Use of Marker Information to Maintain Variability
in Small Populations
H. Takeda, T. Obata, T. Furukawa, K. Nirasawa and H. Takahashi
NewEndocrinological
Methods for Production of Germ Cells in Birds
S. Ishii, S. Wakabayashi
and M. Kikuchi
Use of Chimeric Animals to Preserve Animal Genetic Resources
A. Onishi
Discussion

167
175
183
1 89

Session 5: Use and information
management
Conservation and Use of the Kacang Goat in Indonesia
Survey and Development of Experimental
Information

Discussion

System for Animal Genetic

M.Astuti

193

K. Fukuta
Resources in the Asian Region

203

Y. Yamada and D. E. Steane

215

Animals227

Paper presented at the MAFF Seminar
Nepalese Animal Genetic Resources.
N.P. Shrestha
General229

Discussion
243

Closing

Remarks
H. Seko 247

List of Participants251

INTRODUCTION
Preliminary

addresses

Keynote addresses

Chairperson
S. Muramatsu

Welcome address
TOSHIAKI

HOSODA

Research Councillor,

Council's

Agriculture,

Forestryand Fisheries

Ministryof Agriculture,

Forestry

Secretariat,
Research Council
and Fisheries

1. Distinguishedguests, ladies and gentlemen, on behalf of the Agriculture, Forestry and
Fisheries Research Council, it is my great pleasure to extend sincere greetings and best
wishes to all participantsin this"MAFF International
Workshop on Genetic
Resources ".
Conservation of genetic resources is of great concern with respect to agriculturalresearch,
because genetic resources furnish the basic material for research and is thefoundation of all breeding activities.
This is why many national and international
research centers have genebanks, and are extending their genetic resources research
programs. Worldwide efforts are being made to conserve and use genetic resources ina sustainable way.
Japan, as a member of international
community, will continue to make every
effort in the field of sustainable conservation and use of genetic resources.
2. In addition to our effort of transferring technologies relating to genetic resources,
Japan shares with the international
community the responsibility
for creating acoordinatedglobal genetic resources network. Japan is actively engaged in thediscussions of many international fora. We are a member of the "Commission on Plant
Genetic Resources" in the Food and Agriculture Organization of the United Nations, andalso Japan is a member of the Contracting Parties to the "Convention on Biological
Diversity". Recently, I attended the second meeting of the Conference of the Parties tothe "Convention on Biological
Diversity"
held in Jacarta, Indonesia, and discussed
intensively
with our partners how to realize the spirit of the Convention.
3. Animal genetic resources, the theme of this workshop, comprise an importantcomponent of agricultural
genetic resources. Consequently Japan supported the FAO's
resolution at its 28th Conference to strengthen its activities
on animal genetic resources.
Even before that decision,
Japan had been supporting the FAO's field project

"Conservation and Use of Animal Genetic Resources in Asia and the Pacificll since its
inception in 1992, not only with financial support, but also by dispatchinga specialisttothe project. We fully hope that the recent decision of the FAO Conference will mark thefirst milestone in the worldwide effort to sustainably conserve and use animal genetic
resources.
4. MAFF International Workshop on Genetic Resources is held in part as an expressionof Japan's committment to contribute actively to the conservation and use of genetic
resources globally.
The purpose of it is to enhance international
collaboration
andcooperation and to increase understanding and awareness of research progress among
concerned scientists
in this vital field. Like the previous Workshops, Plant andMicroorganisms Genetic Resources , this third Workshop will offer a chance of sharing
information and experiences on Animal Genetic Resources.
I would like to conclude my address by expressing my sincere desire that thisWorkshop will strengthen our mutual understanding and develop warm and lasting
friendships,
so that it will bring us a solid basis for cooperation in the future.
Thank you very much.

Opening address
MASAHIRO NAKAGAHRA
Director

General, National

InstituteKannondai 2-1 -2, Tsukuba, Ibarakiof Agrobiological

Resources

305, Japan

It is a great pleasure for me to extend a warmwelcome to all participantsof thisworkshop. I would like to particularly
welcome those participantsthat have travelled along way to be with us. I think Dr Bodo has travelled the furthest, from Hungary, andI hope he has overcome his jet lag. We do appreciate you all for sparing the time to joinin this workshop.
I would like to acknowledge and thank the Agriculture, Forestry and Fisheries
Research Council and our sister institutes
who have helped to make this workshop
possible.
Conservation of genetic resources is of central concern globally and seeking
ways to improve global food and basic commodity security for all mankind is among themost urgent tasks facing humanity. All countries are interdependentwhen it comes toconserving, evaluating and using genetic resources. The speed with which the many
issues related to biodiversity
are becoming international
issues involving government
level meetings has implications
for science and scientists.
To help ensure barriers to
germplasm exchange are not erected and cooperation on issues related to germplasm
conservation are enhanced personal contacts and mutual trust and friendship are essential.
Fostering international cooperation and interaction is one of the results which I hope will
emerge fromthe next few days you all have together in Tsukuba.
Animal genetic resources conservation is the particular focus of your discussions
over the next few days. The papers to be presented in this meeting cover a very wide
range of species from the small silk worm to the buffalo. Birds, mammals and insects
will be discussed in different sections of the workshop. Each species has its own
particular characteristics.
Conservation systems for each species differs. However , overthe next few days I hope all participants
will be stimulated by the diverse reports
presented. I hope you will:a) find differences among research which will be of interest in the research you
are engaged in;b) seek commonalities which will make the broad management of conserving
animal genetic resources more efficient.

I would like to take this opportunity
to tell you a little about the National Institute ofAgrobiological
Resources (NIAR) which you will learn more about this week. NIAR has
5 main research sections:
Genetic Resources,
Molecular Biology,
Cell Biology,
Applied Physiology,
Radiation Breeding.
Perhaps I can explain the connection between these sections by saying that each aims toprovide a foundation for improving agriculture. Conservation of genetic resources, aswell as, basic research on molecular biology are essential building blocks in agricultural
improvement. Weare making full use of the advances in biotechnology
and information
sciences to make progress.
Currently our institute is reorganising and extending its research facilities
so thatit will be able to carry out a strong research program into the next century. One of theadditions next year to our facilities will be a new genebank for animal and microorganism
genetic resources.
However, an institute does not stand alone. All our research depends onpartnership both within Japan and abroad. The collaborative linkages we are building andhopefully strengthening
by joint research are an important component of our institutes
vitality.
We are looking forward to enhanced collaborationin Japan and, particularly,
with colleagues from overseas in the future.
Whenweproduce the proceedings of this workshop the questions are faithfullyrecorded. Questions are valuable to seek clarity and stimulate ideas. I encourage allparticipants,
mindful that the proceedings will go to a much wider audience, to make useof the question sessions fully.
Finally, let me wish you an enjoyable time in Tsukuba, so that you may takehappy memories back to your homes, to share with your families over the holiday season.
Thank you very much.

Keynote addressI

Why and Howto Conserve DomesticatedResources

Animal Genetic

IMURE BODO
Department of Animal Husbandry,
PO.Box 2, H-1400 Budapest,

Universityof Veterinary

Science

Hungary

Summary
The organized activity
in recent decades for cultural
The ancientfor maintenanceand technicalanimal breeds are products

There is also the aesthetic
tourism must not neglectedanimal genetic

resources has acceleratedof human selection

and they are importantargument that they are elements of nature and landscapeprotection.

in teaching.
Their role ineither.

The future requirements
be used in harsh environments,

for animal production

are unknown. Non commercial animal breeds canand also in cross breeding

Animals world wide play an important
Conservationof domesticatedreasons.

role in organic

programmes can be organized

as gene resources,

and as control populations.

matter production.
"in situ" or "ex situ" using

traditionaland modern

methods.

Introduction
In the second half of the 20th century the improvement of animal breeds has
accelerated. As a consequence somebreeds have increased worldwide while other noncommercial ones have decreased or even become extinct. The idea of the maintenanceof valuable old things, "antiques", is not a new feature of mankind. In the first place oldmonuments were protected and the concept of protecting wildlife is not new either.
Domesticated animals seem to have been neglected perhaps because they are the productsof human selection and not considered "natural" in the way wild animals are. On theother hand, living beings cannot be compared to buildings
or other man mademonuments.
Only during recent decades has the idea of conservation of animal genetic
resources become increasingly a part of animal breeding, both in theory and practice.
Conservation means the management for human use of livestock so, "that it
may yield the greatest sustainable benefit to present generations while maintaining its

potential

to meet the needs and aspirationsof future generations" (IUCN definition).
According to this definition, conservation touches animal science and husbandry
as a whole, because without this attitude the future genetic diversityof livestock for
mankind would be rapidly lost.
Preservation has a narrower meaning according to the FAO definition which is :"That aspect of conservation, by which a sample of animal genetic resources population
is designed to an isolated process of maintenance, by providing an environment, free fromthe human interference which might bring about genetic change. The process may be "insitu", whereby the sample consists of live animals in a natural environment, or it may be"ex situ", whereby the sample is placed, for example, in cryogenic storage"
The purpose of preservation is to maintain farm animal populations
without
genetic change as far as possible.
The conservation of such domesticatedanimal breeds needs financial support.
Therefore the case for their conservation must be made very strongly (Bodo et al 1 984).
Arguments
There are two kinds of arguments for the preservation
farm animals :* cultural reasons and* technical arguments.
Culturalof genetic resourcesofreasons** The ancient, non commercial breeds are the product of human selection andeffort therefore like old monuments or buildings,
they also deserve protection. Also
domesticated animal breeds are a valuable part of human culture.
** Among these old breeds there are many animals with an attractive
conformation. So their maintenance is reasonable from an aesthetic point of view, Many
artist, painters, photographersand sculptors would understand this arguement.
** Ancient domesticated animal breeds can be used in teaching history andanimal breeding.
They belong to both the history of human life and to the history ofanimal husb andry.
** A landscape with pastures is coloured with the grazing animals and so thebreed is important for the picture of the landscape.
If the breed changes, it involves

changing the environment as well. Thus, if there are countryside protection areas breedsof grazing animals are also important.
** Humanart uses animals as subjects too. For example, the shepherds andherdsmen used to carve artisticallythe long horns of old Hungarian Grey cattle. Also thetools they use are decorated with the pictures of these old fashioned animals. So theancient animal breeds and folk art are well connected.
** The people from the towns highly appreciate the unusual view of these oldfarm animals. Therefore the possible role of ancient, non commercial farm animal breeds
in rural tourism should not be neglected.
It is a potential
source of revenue forconservation of domesticated animal genetic resources.
Technical

arguments** Wedo not know the future requirements of mankind for animal production.
It is possible,that some neglected products of today can come into fashion again.
** There are marginal areas with very harsh conditions for animal production.
Some ancient animal breeds are well adapted to these areas. They are resistant to thediseases and well adapted to climatic conditions.
In some cases in these areas only local
breeds can be used.
** Ancient local breeds are well adapted to harsh conditions and are very usefulin cross breeding systems. In most cases these local breeds are suitable for female lines
and they should be crossed by a terminal male line. The results of such crossing are
economically advantageous if the breeds are well selected.
** The genes of ancient breeds are very important gene resources. The use ofdesirable
genes from these animals is not easy, because they are linked to undesirable
genes. In the future, biotechnology and genetic engineering will help break these linkages
or provide new means of transferring desirable single or quantitative
genes.
** In order to compare the genetic progress in production or in other traits, oldbreeds can have the role of a control population (Crawford, 1 989).
** It is not true, that only native, local breeds can be used in organic production,
but the old fashioned conformation and the good adaptation and resistance of the oldbreeds make them suitable for this use, at least as trade marks of given special products.

10

The most important methods of conservation
The two basic methods are "in situ" and "ex situ" methods.
In situ conservation means the maintenance of the population in its native environment
under traditional
management conditions.
Ex situ conservation includes cryogenic
storage of breeding material (ova, sperm, embryo). Specialists
used to compare these two
methods from the aspect of their relative costs. The general opinion is, that themaintenance of living herds is in most cases more expensive than the storage of deep
frozen material. Thus, there are many arguments in favor of both methods.
Comparison of the maintenance of living herds and cryogenic storage
The most important advantage of cryogenic storage is the lack of genetic change,
if the sampling was quite correct when stored. One can imagine an unexpected effect of,for example, back ground radiation but up to nowduring the first 40 years of deep frozen
storage of sperm in Cambridge no such signs have been observed. In the case of living
animals selection can not be avoided. The changes by selection can be minimized, but
it is impossible
to avoid it totally.
It is nearly impossible to compare the expenses of these two methods. The roughcosts of cryogenic storage are less. Nobody can make, however, calculate thecompensation received from rare breed products sold in the market. The prices of theseproducts depends on the market and it seems there is a tendency for there to be anincreased appreciation
for these products in different markets. It is questionable,
however, whether lower productivityof these breeds can compensated by the higher
prices of special products in the market. The possible income to breeders from thetourism can not be neglected either. Therefore a comparison of costs can not becalculated globally, only in individual
cases.
The advantages of the in situ preservation are that the useful traits of the animals
are not forgotten, and breeders can eliminate some genetic defects, if there are any.
For the in situ herds there is some danger of natural accidents and diseases,
however an evolution of resistance can be supposed against the developing bacteria,
which does not happen in deep frozen situation. The most important source of danger forcryogenic storage is human negligence.
The conclusion of this comparison is that the best means of conservation is to
use these two methods together, the one method can complement the other one very well.

ll

Howto selectpopulations
to bepreserved?
The problem is the choice of the animal populations
for preservation which is
subsidized by the government or club (e.g. the Rare Breeds Survival Trust in the United
Kingdom). Classificationis needed between candidates.
There are many methods in thisrespect such as the Red data book, Alderson (1989),
Hanson (1990),
Bodo (1990b),
andthe North American system (Bixby et al , 1994). I want to mention here only the mostimportant aspects.
1st step.

We have to take into consideration
the grade ofendangeredness
i.e. the risk
status of the given population(see the discussion below on the minimum
number). The smaller the population size the faster conservation measures
should be taken.
We should weigh the pros and cons for the task of a given country topreserve given population
or not. The following questions
should be
answered:* Is the given breed native to the country?
* If not, is it unique (i.e. if in the native country it is already extinct, are there
any other countries where an important population of the given genotype canbe found)?
* Are there close relatives somewhere?
* Has the breed some national merit (belonging
to a specific landscape, or
somehabits of people etc.)?
* Is the breed older or younger than 200 years old?
* Is it a pure population or a mixed one? (Sometimes this is a decisive factor
although a mixed cross bred population
can be maintained as well, if it hasother merits !)If the answer is yes to any of these question conservation is necessary.
Ranking between the population involved into preservation
following aspects :-performance-special products-adaptation

based upon the12-resistance against diseases-other traits (morphological,
immunological,-distinctivenessfrom the other breeds-good combining ability.

ethologicaletc.)

Scientificproblems in preservationof domesticatedanimals
Conservation and preservation are of increasing concern to animal scientists.
Scientists
are faced with an array of new research problems. Some of these research
problems include:
Newconcepts and definitions,the pros and cons arguments, decreasing genetic
variability
in different breeds, the problem of inbreeding, the necessary population size(the minimum number of animals within populations to be maintained),
selection in small
populations,the different methods for maintaining living herds and cryogenic stored
genetic material. I will discuss briefly three of these issues.
Minimumnumber of the individuals in preserved populations
Howmanyanimals are needed for preservation programmes, and what is thelimit for an endangered status. Science has not yet provided definite answers to thesequestions.
In the Genetic Diversityof European Livestock Breeds (Simon andBuchenauer, 1993) the use of effective population size (Ne) has been the basis. In theWorld Watch List ofFAO (Loftus and Scherf, 1993) a category system is used, and asimilar one has been elaborated in America (Bixby et ai , 1994).
It is necessary to create a system accepted worldwide for the minimum numberof domestic animal populations to be preserved for three reasons :-It is important to create and keep up to date a World Watch List or
Regional, National lists of domestic animal breeds in order to observe the vulnerability
status and also to access the risk of genetic drift and extinction.
-It is indispensable
to recommend a population size for each populationneeded for preservation programmes in situ or to give advices for sampling a population
to be involved in preservation programmes ex situ.
-It is possible, that the scientific based grade of risk status will serve for abasis of financial support of the populations in question.
In order to avoid genetic drift and the damaging effects of inbreeding as many

13

animals as possible should be kept. On the other hand, to keep the costs low the
population size of non commercial animals must be as small as possible.
Therefore, acompromise should be found.
There are some problems when using Ne alone as a basis for the minimum
population number so scientists have recommended some other criteria besides Ne, forexample, variations in family size, number of herds, existence or lack of cryogenic
storage, trends in population size, crossbreeding,existence of outstanding traits andspecialadaptations(Maijala,
1992),
distribution,
percentage of migration,therelationshipamong males (Simon, 1 994).
Thus Ne must be complemented with other factors to be of practical use. The
category systems (e.g. Bodo, 1990a; Loftus and Scherf, 1993) are not suitable forpractical
use without other considerations.
Therefore, the FAO and ALBC system
incorporates some of these additional factors.
My conclusion is, that the combining of a category system and Ne is the bestsystem for determining the number of individuals
to have in conserved populations. In
the framework of accepted categories of endangered status i.e. between 100 and 1000
females with an adequate Ne population can be maintained with a minimal genetic loss.
Within these figures it is obligatory
to create a special minimum number for each breedor herd. In order to obtain a correct solution, the peculiarities
of a given species, or breed,
the mating system, the genetic variation within the population, the management, effectof the given environment, the possible generation interval, some practical aspects of herdsize have to be taken into consideration.
The number of breeding animals, the sex ratio are only preconditions
to correctpreservation because the most important point is the replacement. The population age
and sex structure must be maintained in its optimum proportions over time.
Recommendation for a correct replacement system
In the literature there are many opinions concerning the minimum number ofbreeding animals and their sex ratio (Ne) but the most important point, replacement,
also a correct Ne should include conservation of replacement, is scarcely mentioned,
therefore the most important aspects should be mentioned here:-Each breeding animals must be substitutedby his/her offspring.
-The male lines and female families must be maintained.

14-The frequency of all possible

traits(inc. immunogenetic

ones) must not bechanged.
-Generation interval should be adequate for a given breed.
-Priorities must not be given to a favorite progeny group in replacement.
-For higher level of safety reserve animals should be kept.
-The most important precondition for correct replacement in preserved herds isthe retention of the genetic diversity of the parent population in the progeny using many
parental males (narrow sex ratio, high Ne in the preceeding matings).
Yamada andKimura ( 1 984) discuss the details and consequences of different mating systems
Selection

within the preserved populations
In order to maintain the given gene structure of the preserved populations,
theoreticallyall the forms of selection are prohibited.
In practice, however, it is
impossible.
If we do not keep all the males and females for the new generation it isalready a form of selection. Some random selection is possible for big populationsofsmall animals (King, 1982), but it is more difficult in the case of large animals. Simon(1982)
is of the opinion that in preservation secondary traits are important and theadaptation to environments today will be significant
in the future.
The main rules of selection have already been given, because at the moment of
replacement the final decision for the selection is made.
Use of the preserved livestock
The main purpose of preservation of domestic animal genetic resources is tomaintain valuable genes for the future. In spite of this, it is important to speak about apresent use as well, because it is necessary in order to secure funding. Also there are
already good examples of the practical use of animals considered to be non-commercial.
First of all the use of rare breeds in alternative farming and in organic production
should be mentioned. Alternative farming is possible with modern improved breeds or
hybrids as well, but the ancient breeds are better adapted to the harsh conditions. It is agood idea to use the attractive old breeds as trade marks for organic production. A good
possibility
presents itself in National Parks, where the use of artificial fertilizers
isprohibited and the grazing animals are in harmony with the protected nature. The marketof organic product is increasing and becoming more organized in Europe and North

15

America. Certificates fromthe IFOAM (International
Federation for Organic Agricultural
Movements) are given for these products.
Rural tourism can provide some financial help for the maintenance of rare
breeds. Ancient herds can afford an attraction for visitors. Special products are more andmore required in the markets of developed countries e.g. the fat produced by the
Hungarian Mangalica pig breed was neglected because of the demand for lean pork until
recently. Nowa new market is open for old fashioned ham which is well produced bythis ancient breed.
Successful cross breeding experiments have also occurred. The primitive
Hungarian Grey cattle was crossed with Simmental and the Fl with a terminal breed
Charolais gave good economic results compared to other cross bred animals. The use ofcrossing draught horses (Irish Draught, Cleveland Bay or Hungarian Noniusz) withEnglish Thoroughbred stallions give good horses for sporting (Show jumping, dressage
and three day events) purposes.
Conclusion
The conservation and preservation of domestic animals is of increasing concern
among scientists involved in animal science and animal breeding.
The FAO category system is suitable for different Watch Lists. For individual
cases, however, a special population size must be determined, all the specific conditions
taken into consideration.
The maintenance of threatened populations
is possible within the category"endangered", if the scientifically
based methods are used.
In somecases, the use of ancient breeds has already resulted in special products
being produced.
The obligationof governments is to protect and maintain populations of allnational domestic animal breeds threatened by extinction.
References
Anderson,L. 1 898.

The chance to survive.

7LF.143.p
Bixby, D.E., Christman,Livestock
Bodo, I. 1990a.

A.H. Jolly LTD. Yelvertoft

C.J., Ehrman, C.J., Sponenberg,

D.P. 1994.

Manor, Northamptonshire

NN6

TakingStock. (The North American

Census)

McDonald and Woodword Blackburg,

Virginia

1 82 p.

Specialproblems

livestock.

4th WCGALP Edinburghof conservationof domestic

horsesKeynote address II
History and Phylogeny of Japanese
Strategies
for Their Effective Use

Native Animals and

YUICHI TANABE
Azabu University
Fuchinobe

1-17-71 , Sagamihara

229, Japan

Summary
Nine animal species native to Japan or domesticated in Japan are conserved. About 1 0,000 years
ago, the first migrants to Japan (the Jomonese) brought dogs to Japan. About 2,000 years ago, the Yayoi
migrants

brought silkworms,

chickens

have already become extinct.

and the other type of dogs to Japan.

In the Ko fun period (1700-1400

They also brought pigs which

BP), the new migrants

brought

horses andcattle.
In the Heian period(
Kamakura period (1200-1270
China.

were introduced.

In theAD) or the Muromachi period (1 370-1 394 AD), ducks were introduced

800-1 190 AD), cats and new varieties

fromGoats were introduced

domesticatedin the limitedof chickens

parts of the West Japan after 1500

in Japan 400-600 years ago from the migrating

AD. Quail were

wild quail. Many associations

for conservationof the native animal breeds have been established.
Many Japanese native animal species
quality.

Japanese

beef cattle (the Japaneseworms are famous for producing

highsuch as beef cattle,

Black)
qualitysilkworms

and quail have excellent

are famous for their well-marbled

meat. Japanese silkand quantity

quail have high egg

of silk fiber.

Japanese

production

and are used as experimental animals in avian physiology
and genetics worldwide.
Other animal species such as chickens, horses, goats, dogs and cats are kept for their specificcharacters.

Japanese native chickens

consist of 1 7 breeds,

long-tailed

fowl breeds (the Onagadori),

and bantam breeds (the Chabo),the Taromaru and the Koeyoshi) and for delicious
Hinaidori).

Japanese native horses consist

the Hokkaido

is used as

and they have beautiful

meat productionof eight breeds,

and kept for various special

native goat breeds (the Shiba Goat) are consideredJapanese

native dogs consist

They are famous for their simple
Biotechnologies

features.

featuresuseful as experimental

Recently,
animals forof six breeds,

and are becoming popular as companion

and fidelity

to theirowners. Japanese native cats are

They are also companion animals.

for keeping and multiplying

transfer, and cloning and multiplicationof fertilizedof the Japanese native animal genetic

resources.

transgenic

uses. For example,

to carry burdens on rough and mountainous roads in Hokkaido.

small sized Japanese

famous for their gracefulfor their longer crows (the Totenko,

such as game breeds (the Shamo and theruminant physiology.
animals.

external features such asanimals encourages us in adopting

animals with the aid of artificial
embryos, are now essential

Further,

improvements

new strategies

in genetic

for animal breeding

insemination,techniques

embryofor effectiveengineeringuse

to produce

such as to transfer of the

18genes controlling

the resistanceto specificdiseasesarid or humid)

occur in the native animals.

and the resistanceof the stressed

environments(cold,

hot,

Introduction
Recent knowledge concerning the genetic constitution and evolution of animals
and plants helps in understanding
the importance of research in relation to conservationof native animal genetic resources.
Advances in biotechnology,
includinggene
manipulation, in recent years has further increased the importance of conserving native
animal genetic resources.
Eight animal species are native to Japan, dogs, chickens, silkworms, cattle,
horses, cats, ducks and goats. One species was domesticated in Japan, the quail.
Introduced species, including the pig, were brought to Japan from mainland Asia. Native
Japanese pigs have become extinct. Three Japanese native animal species beef cattle,
silkworms and quail have excellent quality.
The other six animal species, chickens,ducks, horses, goats, dogs and cats are kept for their specific characters. An outline of
the history and phylogeny and the present status of these animals will be discussed in thisreview. A list of major domesticated animals in Japan and the time of their migration to
Japan is given (Table 1).
Historyl.Dogsand Present Status

Six Japanese native breeds were assigned as Japanese natural monuments,these
are the Hokkaido(theAinu) dog, the Akita dog, the Kai dog, the Shikoku dog and the
Shiba dog. Two other native dog breeds, the Mikawa dog and the Ryukyu dog are kept
in Tokushima Prefecture
and Okinawa Prefecture,
respectively.
Phylogenetic
relationshipsof Japanese native dogs and native dogs breeds or populations in Korea,
Sakhalin, Taiwan, Indonesia, Bangladesh and Mongolia, and European dog breeds werestudied by using protein polymorphisms
detected by electrophoresisand chemical
analysis (Tanabe, 1991; Tanabe et al, 1991a; Tanabe and Hayashi, 1995). A scattereddiagram of the 52 dog breeds or populations on the basis of the 1st and 2nd principal
component scores is given (Figure 1). It shows evidence of gene flow to Japan in two
directions. One is from Southeast Asia through the Ryukyu Islands to the main islandsof Japan excluding Hokkaido. The other is from Northeast Asia through the KoreanTable1. A list of major domesticatedanimal species

in Japan and the time of migration* Wild mouse were possible** Domesticatedin Japan.

introduced

to Japan.

twice in Jomon and Yayoi periods,respectively.

Table 2. Gene frequencies
wolvesof hemoglobin(Hb) and gangliosidemonoxygenase (Gmo) variantsinpeninsula to the three Japan mainland islands of Honshu, Shikoku and Kyushu. This
studies show that the two genes, Hemoglobin A (HbA) and ganglioside
mono-oxygenase(G/wog), are present only in East Asian dog breeds and populations including Japanesenative dog breeds and populations on the three islands. Interestingly,
HbA and Gmo&is
found in Mongolian wolves {Canis lupus charted) but not found either in European
wolves {Canis lupus lupus) or the Indian wolves {Canis lupus pallipes)(Table 2)(Tanabe and Hayashi, 1995).
These results suggest that dogs were domesticated fromthe West Asiatic wolves {Canis lupus pallipes or Canis lupus arabs), and migrated toEast Asia and they possibly
hybridized
with the East Asiatic wolves {Canis lupus
chanco). Nine million dogs are present in Japan and are kept mainly as companion (pet)animals. Japanese dogs are especially
famous for their fidelity to the owners. Cranialstudies of Japanese people (Hanihara, 1991) and our studies on dogs (Tanabe, 1991)
support the following assumption that the Jomonese brought an ancient type of dog fromsoutheast Asia about 10,000 years ago, and the Yayoi migrants brought a new type ofdog through the Korean peninsula about 2,000 years ago.
2. Chickens
Seventeen Japanese native fowl breeds are assigned as natural monuments ofJapan, these are the Onagadori (the Long Tailed Fowl) the Totenko, the Uzura-chabo, theMinohiki-chabo, the Koeyoshi, the Tomaru, the Minohiki, the Jidori (the Tosajidori,theGifujidori,
the Miejidori
and the Iwatejidori),
the Shokoku, the Shamo(the JapaneseGames), the Chabo (the Japanese Bantam), the Hinaidori, the Ukokkei (the Silkie), the21Kawachiyakko, the Satumadori, the Jittoko (the Creeper) and the Kurokashiwa. Other
native breeds the Chan (the Utaichan), and the Nagoya (the Nagoya-cochin) are kept in
Okinawa Prefecture and in Aichi Prefecture, respectively.
Most of them are famous fortheir beautiful
external features such as the long tailed breeds (the Onagadori, theTotenko and the Shokoku), bantam breeds (the Chabos) and for their long crows, up to
10-20 seconds, such as the Totenko, the Tomaru , and the Koeyoshi. The length of tailof the Onagadori can reach more than 10 meters. Some of them such as the Shamo andthe Hinaidori produce high quality meat. In recent years, a hybrid (the Tokyo Shamo)
between the Shamo males and the Nagoya females are used for broiler production in
Japan, its meat is of higher quality than previous broilers, which were a hybrid between
the White Cornish males and the White Plymouth Rock females (Oishi, 1992).
Phylogenetic relationships
between 20 native fowl breeds or strains in Japan andadjacent areas and five exotic (European and American) breeds or strains, the White
Leghorns, the White Plymouth Rock, the Rhode Island Red and the White Cornish arestudied using blood protein polymorphisms
and the results are shown (Figure 2).
Relatively
close relationships
are found among most of Japanese native breeds,
suggesting that the Totenko may be the ancestor of the Onagadori (Tanabe et al , 1 99 lb).
The oldest bones of chicken have been found in the Harunotsuji Remains (2,000BP) on Iki island, between Japan and the Korean peninsula. The first chicken are thought
to have been brought by the Yayoi migrants about 2,000 years ago.
3. Silkworms
The silkworm moth (Bombyx mori) was domesticated for production of silk fiberfrom Bombyx mandarina in China about 5000 years ago. Tetravoline silkworms wereintroduced to Kyushu, Japan, from middle China, and trivoline silkworms from Korean
peninsula to Kyushu in the early and middle Yayoi period, respectively.
This has been
deduced because the diameter of silk fiber found in Yoshinogari and Asahikita Remains
fromthe middle Yayoi period, is finer than those found in the remains of the early Yayoi
period (Nunome, 1988). Productivity of silkworms in Japan is very high using F i hybrids
and other excellent breeding systems including genetic engineering techniques (Taj ima,
1984).

Figure1. Relative position of 52 dog breeds or populations
defined by the first (Zl) and the second (Z2)largest principal components of the distance matrix based on variance-covariance analysis of genefrequency at 16 polymorphic
loci. Figures in parentheses
represent the contribution
to the total
variation (Tanabe and Hayashi, 1 995).

Figure2. A dendrogram of the genetic distance matrix computed from the gene frequencies of 1 7 variable
and non-variable loci using Nei's equation among 1 9 Japanese Native fowl breeds, five Europeanand American fowl breeds and one Korean fowl breed, the Chejudo native fowl (Tanabe et al ,1991b).

Figure3. A dendrogram drawn from genetic distancesamong the Asian and European horse populations.
S, S-M and M means small-sized,
small to medium-sized,
and medium-sizedhorses, respectively(Nozawa e* <i/., 1984).

24

4.PigsPigs were domesticated fromwild boar (Sus scrofa) in China about 1 1 ,000 years
ago, possibly other types of Sus scrofa were domesticated in the West Asia about 9,000
years ago. The Yayoi migrants brought pigs with them about 2,000 years ago. Japanesepigs were not thought to be descendants of Japanese wild boar, because the domesticated
pig skull especiallyteeth were much bigger than those of the wild boar of Japan at theperiod. The native wild pigs became extinct in the beginning of the historic age (shortly
after the introductionof Buddhism, around 600AD) in Japan.
5. Cattle
European cattle breeds (Bos taurus) were domesticated from aurochs (Bosprimigenius)
about 9,000 years ago in West Asia. Two Japanese pure native cattle
populationsin Mishima, Yamaguchi Prefecture and in Kuchinoshima, Kagoshima
Prefecture, are descendants of aurochs (Bos primigenius),
not the Zebu (Bos indicus).
European-Korean-Japanese breeds or Japanese native populations have a much higher
incidence of HbbA (A type of hemoglobin B) than HbbB, while HbbB are predominant
in zebu-type cattle (Namikawa et al , 1984). The shape of the Y chromosome of cattle
descended from aurochs including Japanese pure native cattle, is sub-metacentric and thatof the Zebu cattle is acrocentric. Many breeds of Japanese beef cattle have been bred forthe production of high quality meat. Japanese Black cattle (Wagyu) are from hybrids
between Japanese native cattle and the Brown Swiss or the Devon. The Japanese Blackis famous for the production tender meat with a much smaller size of muscle cell fiberand a very high score for marbling compared with the western beef breeds.
6.Horses
Eight Japanese native horse breeds, i.e. Hokkaido, Kiso, Misaki, Noma,
Tsushima, Tokara, Miyako and Yonakuni are conserved in Japan. Phylogenetic
studiesby Nozawaet al (1976) on Asian and European horse breeds and populations suggestthat all of the Japanese native horse breeds are descendants of horses which came throughthe Korean peninsula with migrants in the Ko fun period, and not through Taiwan fromSouthern China. A dendrogram drawn from genetic distances among the Asian andEuropean horse populations
is shown (Figure 3). A significant
difference is observed
between the Asian horse breeds including the Japanese breeds and the European breeds

25

including the Arab, whereas no consistent trends in body size are observed among thebreeds or populations examined in this study.
7.CatsAll domestic cats were derived fromthe African wild cat, Felis silvestris lybica,and domesticated to destroy rodent pests in Egypt 3,500-4,000
years ago. The first
historical records of cats in Japan appeared in 889AD (the early Heian period). It is said
that Japanese envoys to China in the Tung dynasty brought cats with them to protect
Buddhist scriptures and cocoons of silkworm moths from rats in the ship. More than six
million cats are kept in Japan mainly as companion (pet) animals. Most of them areJapanese native cats which are descendants of the cats introduced
from China
1,200-1,300
years ago. They are famous for their graceful features. Blood protein
polymorphisms in Japanese cats were studied by Nozawa et al (1 985). Recently, many
hybrids between the Japanese native cat and the Siamese or the Persian cat have been
introduced into Japan.
8. Ducks
The duck is an important domestic animal, especiallyin Asia. Ducks have thesecond largest (68 1 million) population among poultry species. Eighty six percent of all
ducks are found in East and Southeast Asia.
The ancestor of the domesticated duck was the mallard (Anas platyrhynchos)which migrates between northern and southern parts of Eurasia.
Duck have been
domesticated in China for at least 3,000 years. It is quite certain that the major center ofdomesticationof ducks is the area from East Asia to Southeast Asia. The earliest
evidence of domestic ducks in Europe is as recent as the 12th century AD, and ducks
were introduced frequently to Europe after 1493 when ships able to navigate to and from
Asia via the Cape of Good Hope (Delacour, 1964). The egg production of ducks in East
Asia especiallyin China is very high and exceeds 300 eggs per year (Clayton, 1984).
Ducks were introduced to Japan in the Kamakura period (1200- 1270) or the
early Muromachi period (1370- 1394 AD) possibly from China.
In Japan, there are three indigenous duck breeds, the Kairyo Osaka, the Aokubi
Ahiru and the Naki Ahiru. The Kairyo Osaka originated from hybrids between theJapanese native white duck and the Pekin, which was imported from USA, and the hybrid

26

wasbackcrossed with the Pekin. Of two flocks of Mallards examined, one has been kept
in a farm in Kyoto Prefecture, Japan, after a recent catch of migrating mallards (Mallard1), and the other may have descended from hybrids between migrating mallards and theJapanese native green headed ducks (the Aokubi Ahiru)(Mallard
2).
Phylogenetic
relationshipsof duck breeds and populations were studied using
two dendrograms drawn by genetic distances among the breeds by Nei's equation. Figure
4 shows a dendrogram based on protein polymorphisms detected by using electrophoresis
on 22 genetic loci (Tanabe et aL, 1988; Tanabe, 1994).
Figure 5 shows anotherdendrogram based on DNApolymorphisms detected by randomly amplified polymorphic
DNA (RAPD) patterns (Okabayashi,
Kawasaki and Tanabe, 1995). The dendrograms
show similar results.
These results indicate that a marked difference of genetic constitution
is present
between the duck breeds in Southeast Asia and those of Northeast Asia, indicating thatancestral mallards are different between the two areas. It is plausiblethat the first site fordomestication of ducks was China, and the second site for domestication of the species
was Southeast Asia including Indonesia and Malaysia.
A close relationshipbetween the Khaki Campbell and most of the Indonesianduck breeds except those fromEast Indonesia is consistent with the report that the Indian
Runner was the one of the founders of the breed. It is possible that European breeds werefirst introduced from Southeast Asian ducks and or Chinese ducks, and they were matedwith European mallards later.
These present studies also shows a similarity among the Northeast Asian ducks
including the Kairyo Osaka, a Japanese breed and the mallard (migrating mallards).
Itis postulated that Japanese native duck breeds are descendants of the hybrids between the
Chinese ducks, which were introduced into Japan 700-800 years ago, and mallards, whichwere migrating between Siberia and Japan.
9. Goats
Goats were domesticated mainly from the Bezoar goat {Capra aegagrus) inWest Asia. Goats are very important animals for meat production in Asia, and adapted
to both hot and arid or hot and humid climates. Goats were first introduced to Japanthrough the Ryukyu (Okinawa) islands after 1,500 AD. Japanese native goats are foundin Okinawa Prefecture (the Okinawa goat), Tokara islands (the Tokara goat) and the27

Goto islands and interior of Nagasaki Prefecture (the Shiba goat). These goats are small
and are useful experimental animals for ruminant physiology.
Phylogenetic
studies on goat populations in Asia show the close relationships
amongthe Okinawa, the Shiba, native goats in Korea and Taiwan, and Japanese Saanen(an exotic breed) and among the goats in Philippines,
Indonesia (the Etawa and localgoat) and Bangladesh (the Black Bengal and the Jamunapari). A dendrogram based oncluster analysis of goat population data is shown (Figure 6) (Nozawa and Katsumata,1984).
10.Quail
The Japanese quail (Coturnixjaponica)
was domesticated in the Muromachi
period or the early Edo period (400-600 years ago) in Japan. It is the only animal species
to have been domesticated in Japan. Egg production rate of the species has improved
since the begining of the Meiji era. Wild quail still inhabit Japan and migrate between
Siberia or Sakhalin and mountain regions of Honshu in Japan. Kawahara( 1976) caught
306 wild quail at 800-850m in grassland at the foot of Mt. Fuji from October toNovember between 1965- 1968, and bred them in captivity.
Their egg production rateduring the first 60 days was 44% and age at the first laying was 1 10 days. After 10
generations of domestic propagation without conscious selection, the rate increased to79%, 90% of the level of the domesticated quail. The age of first laying decreased to
60 days compared to 49 days for the domesticated quail (Figure 7). While the coefficientof variation of these traits decreased somewhat during the 1 0 generations, much greater
variation for the traits was observed in the wild quail and semi-domesticatedones
compared with domesticatedquail (Figure 8). These results strongly suggest that thecontribution to the improvement of productivity
during the domesticationof the species
was due to recombination of pre-existing genes rather than due to mutations. The egg
production in captivity of wild ancestral species of domesticatedbirds is high in suchbirds as mallards and Japanese quail, although the domesticated birds always produce
more eggs than their ancestral wild ones. It is thought that the high productivityofdomesticated birds is an expression of natural fecundity given appropriate opportunityand owes little to artificial
selection.
In the wild, the expression of genetic variability
would not have been possible by natural selection and makes it possible for animals andbirds to adapt the new environment. The interventionof man introduces drastic

Figure4. A dendrogram of the genetic distance matrix computed from the gene frequencies of22 variable
and non-variable loci using Nei's equation among 16 duck breeds and strains, and two mallard
flocks (Tanabe, 1 994).

Figure5. A dendrogram of genetic distance matrix computed from gene frequencies
polymorphic
DNA (RAPD) among 16 duck breeds and populations(Okabayashi,
Kawasaki and Tanabe, 1 995).

of random amplified
and a mallard flockFigure 6. A dendrogram showing genetic distance of 1 1 goat populations.
belong to the Indonesian populations,and the Black Bengal
Bangladash-India
breeds ( Nozawa and Katsumata, 1 984).

The Katjang , the Etawa and local
and the Jammnapari belong to the

Figure 7.
Changesin
egg
production(O)andsexual maturity (•E) of
wild
Japanesequail
during 1 0 generations in
captivity
without
conscious
selection.
Verticalline represents
standard
deviation(Kawahara, 1 976).

Figure 8.
ChangesinsexualmaturityofwildJapanese quail during 8
generations
in captivity
without
conscious
selection(Kawahara,1976).

Figure9. A scattered plot of quail populations(Kimura, 1994).
O Japanese commercial strains; ©Wild quail population
caught in Japan,
å¡Commercial strains in Canada and France, å Experimental strains in Canada,
J(UBC-J), G(Giant) and F(FRA2)
are populations
bred for larger sizes.

31

environmental changes. These changes make possible the expression of genetic
variability.
After capture,
reproductive
fecundity
is much improved during
domestication without conscious selections in duck and in the Japanese quail.
Phylogenetic
studies on wild and domestic quail indicate that some genetic
differences are observed among the commercial strains for egg production in Japan, theexperimental
and commercial strains for meat production in Canada and Francecompared to wild quail in Japan (Figure 9, Kimura, 1994). As all the quail strains inthese studies originated from Japanese quail (Coturnixjaponica),
the genetic difference
observed among the groups is derived from genetic drift. Genetic variability
expressed
as average heterozygosity(H) is higher in the commercial strains (0.098)
than theexperimental strains in Japan (1. 189) and in Canada and France (1. 176) and wild quailin Japan (0.078)(Kimuraand Fujii,1989;
Kimura, 1994). The higher variability
observedin the commercial strains may be due to the breeding systems to express effectively
heterosis for a higher egg production. Interestingly,thequail bred for egg production in
Japan weigh 100-200g, and those bred for meat production in Canada and France weigh250-350g. Egg production rate is very high in the commercial strains in Japan reaching
300 eggs per year. Quail are also very useful as experimental animals in avian
physiology
and genetics.
Effective

use of native animals
Recent progress in genetic engineering, which allows us to manipulate genes,
increases the importance of conservation of indigenous breeds or species which are well
adapted to the environment of various Asian regions. For efficient conservation ofgenetic resources, long term storage of frozen sperms and embryos of valuable animals
at -1 96 °C, and the application of artificial insemination, embryo transfer techniques, andcloning and multiplicationof fertilized embryos are very useful for reducing the numbersof animals that need to be maintained. The production of transgenic animals and sexingof sperms or fertilized embryos allow us to find more effective way of using genetic
resources in animals and poultry (Tanabe, 1989). Further improvements in genetic
engineering to produce transgenic animals (Palmiter et al, 1982; Pursel et al, 1989)
encourages us in adopting new strategies for animal breeding, such as, to transfer genes
controlling
the resistance to specific diseases and adaptation resistance to the stressed
environments (cold, hot, arid or humid) which occur in native animals.

32
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areas

35

Questionsand answers in Keynote address session

Q: Referring to the theme of the workshop, how efficientand effective is theconservation of animal genetic resources? (Muladno)
A: Animal genetic resources conservation was initiated only a few years ago. Time isneeded for the system to be effective and results to be obtained.
The leader
organization in this effort is the FAO. In some countries there are good results fromconservation of farm animals, for example the United Kingdom and Hungary?
(Bodo)
Q. Dr. Bodo can you give your ideas on balancing conservation of animal genetic
resources and national programs which encourage crossbreeding
programs?
(Mathius)
A. The two concepts crossbreeding and preservation are not contradictory
ones. Bothare important and complementary. Local adapted breeds properly maintained can be
useful material for crossing with exotics? (Bodo)
Q. Dr. Bodo is the objective of animal conservation the species or the breed.
A. The most important objective is the gene. It can be found in breeds. Species are notthreatened with extinction, as far as domesticated animals are concerned. If weconsider preservation of breeds, it is not the name that is important, but the valuablegenes in them. Therefore, the different varieties within breeds should not be neglected
either.
When wepreserve breeds, species will be conserved automatically,
but we have tolook in the breeds structure (sub-species,
lines, families etc.) in order to save thevaluable genes. (Bodo)
Q. Dr. Bodo, you discussed ex-situ verses in-situ conservation. Between these two
approaches where do zoos fall? (Vaughan)
A. In the strict sense of the word the activity of zoos in conservation should beconsidered "ex-situ" conservation. The role of zoos is very important, for example
the Przewalski horse, and the maintenance of special separated lines of other rare
breeds? (Bodo)
Q. Dr. Tanabe, you showed that worldwide cattle are the most numerous of domesticatedanimals for food. However, Japan has more pigs and much fewer sheep compared to
the world average. Could you explain this? (Ishii)

36

A. Pigs are more efficient than beef cattle in meat production. Most Japanese are lactose
intolerantso cannot drink much milk, unlike Western people. The Japanese
government led by Mr. Toshimichi Ohkubo imported many sheep but most of themdied. Sheep are not resistant to the hot, humid climate of Japan. (Tanabe)
Q. Dr. Tanabe are there still water buffaloes in Japan? (Tan)A. There are a small number of water buffaloes in the Ryukyu islands, less than athousand. They do not appear on the FAO census figures because the number is so
few. (Tanabe)

TECHNICALREPORTS

Session1

Economic evaluation

Chairpersons
H. Mikami
L.V. Ly

Economic Evaluationof Mongolian

Sheep Genetic Resources

TSE SRENJAV ZHANCHIV
Instituteof General and Experimental

Ulaanbaatar-5

Biology,

MongolianAcademy of Sciences

1, Mongolia

Abstract
Mongolia is known to be one of the world's traditionaland selective

breedingof animals

pure breed of animals well adjusted
husbandry employs inexpensive

startedin Mongolia

to the natural
technology.

centers of husbandry.

some five thousands

The domestication

years ago. As a result of this aconditions of Central Asia was developed. The establishedThis experience is an integral part of the world husbandry

heritage.
adjusted

It is possible to use the basic stock of Mongolian
to climatic specifics of Mongolia.
Presently

there are 27 million

livestockin Mongolia,

sheep for selectionof these 14 million

and breed new lines moreare pasture sheep stock.

I. Mongolian Sheep Breeding Lines
For thousands of years Mongolian sheep basic stock have been used forbreeding. It is possible to distinguish
two basic groups of sheep: short fat tail and wide
fat tail. This could be explained by the fact that sheep are use primarily for their meatand fat.
Twogroups could be further divided into several sub-groups :L Khalkha Breed Sheep (Figure 2, 3 and 4)
The largest number of Mongolian sheep belong to this breeding line, They arestrong, and well adapted to Central Asia climatic conditions.
They are well adjusted to
pasture husbandry. Usually they have a dark brown bald head. They are mainly used formeat and fat.
2 Bayad Breed SheepThis breed of Mongolian sheep is raised for meat-fat production. It has a strong
body and usually has white head. There are estimated to be 100,000 sheep of this group.
3. Gobi-Altai Breed SheepThis line is grown mostly in Bugat and Tseel somons (municipalities)
in GobiAltai province (Figure 1). They belong to the fat tail group, and compared to other40

Figure1. Provinces in Mongolia.
1 : Bayan-Olgiy,
2: Uvs, 3: Hovd, 4: Zavkhan, 5:Govi-Altay,
6: AyanHongor, 7:Azhangay,
8: Hovsgol,
9:Bulgan,
10: Ovorhangay,
ll: Selenge,
12: Tov, 13:
Dundgovi, 14: Omnogovi, 15: Govi, 16:Dornod,
17:Suhbaatar
and 18: Dornod. Bar = 150 km.

breeds are characterized by bigger bodies and better quality wool. They are well adapted
to high mountain pastures and weather conditions.
Presently there are about 20,000
heads of this line.

4. Darkhad Breed SheepThis line is grown mostly in the three northern somons of Huvsgul province.
Compared to Khalkha lines they belong to wide fat tail group and are adapted tomountain pastures.
5. Uzemchin Breed SheepThis line belongs to the short fat tail group. Compared to Khalkha line they have
similar body type but can gain weight in a short time. In the first year they have 60
percent of the weight that a grown up sheep hase and in the second year they already have
87 per cent. They are grown mostly for meat. They are well adapted to open steppe
pasture conditions.

FigureFigure3. Khalkha

sheep line (male).

2. Flock of Khalkha

sheep line.

Figure4. Khalkha

sheep line (female).

42

Table

1.

Mongoliansheep body weight

B re e d s an d lin e s

and wool output

B o d y w eig h t (k g )W o o l o u tp u t (k g )K h a lk a h

60

47

1 .4

1 .1

B ayad

73

55

2 .0

1 .6

G o b i‑A lta i

58

47

2 .3

1 .6

D a rk h a d

69

51

1 .8

1 .6

U z e m ch in

74

53

1 .3

1 .0

B a rg a

62

50

1 .3

1 .1

S a rtu u 1

63

52

2 .0

1 .5

T a b le 2 .

A lle le fre q u e n c y o f T f a n d H b o f M o n g o lia n sh e e p
G e n e fre q u e n c y

G ene

A lle le s

H b

Tf

M o u(mn ta= in
3 2‑step6) pe

S e p p(me p=o4p6u5latio) n

D e s ert
(m p= o6 p6 u8latio)nA

0 .2 12 ｱ 0 .0 3

0 .2 1 0 ｱ 0 .0 3

0 .1 4 6 ｱ 0 .0 2

B

0 .7 8 8 ｱ 0 .0 3

0 .7 9 0 ｱ 0 .0 4

0 .8 5 4 ｱ 0 .0 2

D

00.3
.2.2 00264 ｱｱ 00 .0.002230 .14.2 70 091 ｱ 0 .0 43.30 .1.3 652 31 ｱ 0 .0 21

M

00 ..1
0 279 40 ｱ 0 .0 821

0 .0.2 07 90 ｱ 0 .0
.0 023 1

0 .0.3 05 473 ｱ 0 .0 210 2

6. Barga Breed Sheep
Barga Breed sheep are well adapted to steppe pastures, and have a little bigger
and longer body than
Khalkha sheep.‑ Presently there are
140,000 heads
in this line
H‑0 .0 0 1 ｱ 0 .0 0 1
developed since 1 980 in Khulunbuir somon of Dornod province (Figure 1).
7. Sartuul

Breed SheepThis breed has a large strong body and are well adapted to highland pastures.
They usually have a black and brown, bald head and are grown primarily for their meat.
Presently there are 60,000 heads of this line in Erdenesaihan
somon of Zavkhan province(Figure 1).
II.

Results of Mongolian Sheep Genetic Resources
The genetic research of Mongolian animals is not complete yet for several
reasons. Genetics as a branch of science, compared to other countries, was late to43
Table 3. Hb and TfAllele

freauencyof four sheep lines
A llele fre q u en c y

G ene

A lle le sKn h= a 2lk 7h 3a

U nz em= 2c6h 9in

G no b= i‑A2 5 lta
6 in = 251

H b

A

0 .8
.1 09 81 ｱｱ 0 .02 21

0 .8.1 9100 ｱ 0 .2
.0 02

00 .8
.1 27 28 ｱ 0 .0 2

0 .8
.1 27 19 ｱ 0 .0 11

D

0 .2.1 45 20 ｱ 0 .0
.0 2

0 ..12 65 28 ｱ 0 .0 2

0 .1
.3 08 93 ｱ 0 .0 2

0 .3
.1 81 18 ｱ 00 .0 21 1

J

0 .3 2 2 ｱ 0 .0 2

0 .2 9 4 ｱ 0 .0 1

0 .2 2 2 ｱ 0 .0 1

0 .3 1 9 ｱ 0 .0 2

M

0 .2 62 23 ｱ 0 .0 10 .0
.2 643 245 ｱ 0 .0 201 2

0 .2.0 07 295 ｱ 0 .0 201 2

0 .0.1 61 67 ｱ 00 .0 24

T fdevelop.

Also there is a shortage of research facilities
and equipment.
Given the importance of genetic training specialists
in thisfollowing priorities H :a) study the specifics of Mongolian sheep geneticb) use genetic studies for selection and breedingfield

have theGenetic Specifics of Mongolian Sheep
Presently we have studied Hb and Tf alleles of Mongolian sheep and also haveidentified
some alleles based on wool hair color analysis.
Beginning in 1970 mostresearch was concentrated on a comparative study of the main groups of sheep divided
according to climatic, geographicaland other factors. Table 2 summaries the findings:
Mongolian sheep Tf and Hb structure system does not differ from sheep in othercountries.
However, HbB, TfG, TfG, TfD? TfJ alleles frequencies are higher in
Mongolian sheep.
TfP an TfS, TfH alleles frequencies tend to be rare. TfH allele frequency for thedesert populations is, for example, only 0.00 1. Therefore, based on the hypothesis ofHarrison (1973) that gene frequency variation within 0.001 - 0.005, given sufficientnumbers of animals are tested, means a newmutant allele, we can conclude that this newline may have resulted from hybridizationof Mongolian sheep with other breeds.
There is not much difference in Hb allele frequency and gene structure among
Mongolian sheep groups (Table3).
Note there is a difference in Tf allele.
Wefind some differences between Khalkha groups from the Gobi-Altai line in44

Tfand fromthe Bayad line in TfM, TfP alleles.
Such difference can be explained by thelong period of separate breeding and local conditions.
Mongolian sheep are an inexpensive means of wool and meat production well
adjusted to conditions of a nomadic husbandry based economy. However in terms ofmeat and wool the output is rather low. So there is a need to improve the output by
hybridizationand selection of the wool-oriented line (Orkhon breed) and the meatoriented line (Touguud breed).
Currently breeding of West Mongolia long fat tail sheep with wild mountainsheep (Argali) is being conducted in order to produce a more productive meat line. Also
experiments are being carried out to cross Mongolian sheep with karakul sheep.
Another aim of the institute's
research is to increase the number of births per
sheep. Presently about 10 per cent of sheep give birth to two lambs and attempts todevelop a special breed with a greater number of births appears to be successful.
Since 1994 a group of Japanese scholars, headed by Prof. Yuichi Tanabe, hasbeen actively involved in research on Mongolian domesticated animals. When this joint
research project is finished we will have a clearer picture of genetic specifics ofMongolian domesticated animals including sheep groups and lines.

Economic Evaluationof Pig Genetic

Resources in ChinaAIGUO WANG
Collegeof Animal Science and Technology,

Beijing

100094,

Beijing

Agricultural

University

China

Abstract
Pig genetic resources in China are rich and diverse.

This paper discusses

economic evaluationofChinese pigs and analyses their use. The genetic characteristicsof Chinese pigs for prolificacy,meat quality,
stress resistance and miniature body size are reviewed. The excellent reproductive performance of Chinese
pigs (especially

Taihu pig) has been investigatedas an important

project

in many countries.

Chinesepigs arebeing used to study the genetic mechanism of reproduction for improving pig breeds and are being used incrossbreeding
programs for commercial pig production.
Chinese pigs can contribute to improvements infuture pig production

worldwide.

Introduction
Pig production in China has a long history. As early as 9000-6000
years ago,
local wild pigs had been domesticated by the Chinese (Zhang et al, 1990). Pig breeds
diversified over thousands of years of selection and breeding. According to the book "Pigbreeds in China" (Zhang, 1986), there are 48 native pig breeds which can be divided into
6 types and these are a major gene pool for pig breeding worldwide.
2000 years ago, Chinese pigs were introduced to the Roman Empire to improvetheir local pigs and thus bred the Roman pig. In 1770-1780,
Chinese pigs wereintroduced to the UK and crossed with local pigs and the famous Yorkshire and Berkshire
weredeveloped. In 18 16-1817, Chinese pigs were introduced into USA and the PolandChina and the Chester White were bred. Chinese pigs played an important role in thedevelopment of some world famous pig breeds.
In recent years, due to the shortage of pig breed resources, manycountries have
paid great attention to Chinese pigs (especially
the Taihu pig) with high reproductivity.
These have been introduced into many countries to study the genetic mechanisms offertility for improving pig breeds and for use in crossbreeding programs for commercial
pig production, and have achieved a great progress.
Genetic

Characteristicsof Chinese Pigs
Chinese native pig breeds have some unusual characteristics
Western breeds.

compared with the46

1. Prolificacy
Most Chinese native pigs are prolific, except some breeds which belong to theSouth China type, the Southwest type and the Plateau type. The Minzhu has a litter sizeof 13.5 and the Taihu pig of 15.3. There are 7 strains of Taihu pig and they have similar
reproductive performance. Erhualian pigs have the largest total number born and number
born alive. The litter size of Taihu pig from Taihu region is shown in Table 1 (Zhang,1991).

The large litter size of Taihu pig is probably due to the high ovulation rate
(Wang, 1990). The ovulation rate for sows averages 28. 16, and was 6.58 and 6.76 morethan the mean of that of other Chinese native breeds and Western breeds, respectively,
which indicated that farrowing potential is very great in Taihu pig (Zhang, 199 1). And
secondly, many authors (e.g. Bidanel et a/. , 1990; Legault and Bidanel, 1992; Terqui etai,1992;
Xu, 1992) found that the large litter size of Taihu pigs is due to a low
embryonic mortality. The early embryonic mortality rate for Taihu pig and Western pigs
averaged 19.99% and 28.40-30.07%,
respectively,
and that was 8.41-10.08%
lower inTaihu pig than in Western breeds (Chu, 1995).
2. Meat quality
Chinese pigs also have characteristics
of superior meat quality.
According tostudies of meat quality for 10 native breeds, the results indicated that they were superior
in meat color, marbling, muscle fibre diameter, intramuscular fat, dry matter, crude fat,water holding capacity in muscle, flavor, juicinessand tenderness than Western breeds(Chen, 1989)(Table2).
3. Resistance and adaptation
Investigations
on resistance and adaptability
indicate that Chinese pigs have highstress resistance.
In particular,some breeds have cold resistance(Minzhu),heat
resistance (South China type) and adaptation to high altitudes(Tibetan pig and Neijiang)(Xu, 1989; Zhang et al, 1990).
Almost all Chinese pigs are PSS (Porcine Stress Syndrome) free.

47

Table

1.

Littersize of 7 strains of Taihu pig

stra inN o . litte rs

N o . b o rn

N o . b o rn a liv e

E rh u a lia n278

1 5 .9 3

14 .12F e n g jin g

2 20

1 5 .1 7

13 .1 3M e ish a n

170

14 .7 913 .9 0J ia x in g B lac k

2 93

1 5 .0 2

13 .7 3M iz h u82

14 .6 5

13 .3 8S h a to u w u78

14 .3 01 3 .4 9H e n g jin g

93

14 .2 8

1 3 .5 9

T a b le 2 . M e at q u a lity o f n ativ e b re e d s in c o m p a ris o n w ith W e ste rn p ig s l
M P

H LE

JQ H

EH L

JX B

JH

N J

X P

LB W

o p tim al d e n sity

+ 0 .2 8 *‑+ 0 .5 6 *‑‑+ 0 .0 3 6+ 0 .0 9+ 0 .0 3‑

p H v a lu ed rip lo ss %+ 1 .4 0 * + 0 .5 0 * + 0 .5 9 *‑6 .0 4‑‑+ 0 .0 5‑2 .0 9‑‑2 .0 5+ 0 .2 9‑+ 0 .3 3‑4 .8 8+ 0 .0 2‑3 .0 7d Ja m ete r b u e"1 8 3 1"

14 3 1‑4 ‑5 8 ‑1 8 .3 4 * *‑ 19 .8 1* ‑1 8 .3 6 * ‑8 .2 7 ‑1 2 .4 9 ‑ 1 6 .7 1

m s

‑4 0 4 *‑4 3 1 *reb %c ru d e fa t %+ 1 .1 8 + 5 .5 8

"6 ‑8 3‑‑ ‑3 8 *+ 2 .8 6

3 ‑7 4 * "3 8 8 *+ 2 .2 4‑1 0 8 *+ 0 .1 2 + 2 .1 0

D W Z

a v e rag e+ 0 .2 3 * + 0 .2 0‑
+ 0 .4 6 * + 0 .3 2
‑2 .9 4‑
‑3 .8 9"2 2 4 *‑‑ ‑2 8 * " 1 3 6 *"'2 2 6+ 1 .4 5+ 2 .0 1 + 2 .5 7‑

+ 2 .3 2

1 Figures are presented for difference between the Chinese native breed and control breed(Landrace,
Large Whiteand Harbin White); MP=Minzhu, HLE=Hetao Large-ear, JQH=Jiangquhai,
EHL=Erhualian,
JXB=Jiaxing
Black, JH=Jinhua, NJ=Neij iang, XP=Xiang pig, LBW=Large Black-white,
DWZ=Dawuzi.
* P<0.05, ** P<0.01

Table 3. Goal traits
goal trait
litter size
daily gain
feed efficiency
lean meat content
meat quality

and selection

criteria

of Chinese

native pigs
selection

criterion

litter size born alive
average daily gain
feed conversion ratio, average daily gain
lean meat content of carcass, back fat,eye muscle area
meat color, pH value, water holding capacity, intramuscularfat

4. Miniature body size
There are several mini-pig breeds in China. Xiang pig, Bama Xiang pig,
Wuzhishan pig, Banna mini-pig and Small-ear pig of Province Guizhou, Guangxi,
Hainan, Yunnanand Taiwan, respectively.
The body weight of adult pigs is about 40kg.
Chinese mini-pigs have a small body size, slow growth, low metabolic rate, early sexualand body maturity, genie purity, resistance to disease and excellent meat quality. They
are an ideal laboratory animal for medical research and also an ideal raw material for

48

roasting(LianetaL,

1992;

Liu,

1992;

Pei etaL,

1992;

Sung etaL,

1992;

Wang etal,

1992).

These genetic characteristicsof Chinese pigs are valuable and also stable incrossbred offsprings.
Therefore, they could play an important role in improving pig
breeds.
Economic Evaluation of Chinese Pigs
1. Breeding goal for Chinese pigs
The breeding goal for Chinese pigs is to obtain the best possible overall animal
for the food industry with consideration
for future commercial conditions of production.
A definition of the breeding goals includes-economic definition of the breeding goal-the improvement of the animal-considerations
related to production on the commercial level-future needs.
The quantitative
breeding goal for the population is performed in two steps :-choice of the traits in breeding goal
-weighting traits with economic coefficientsor values.
The economically orientated breeding goal leads to establishmentof the total
breeding values which is a linear function of the quantitative
breeding goal. In principle,
all traits influencing efficiency of commercial pig production should be included in the
total breeding values and weighted by their economic values :AT=SWiAi
where Aj is the total breeding value, Wi is the economic weight for trait i, A{ is thebreeding value for trait i.
2. Breeding goal traits and selection criteria
All traits with a major impact on efficiency of commercial pig production should
be included in the breeding objectives. However, the estimation of breeding value is moredifficult and its accuracy and genetic gain of each trait decreases with increasing numberof traits. Therefore the term "majorll implies also that a limitation
should be set to thenumber of traits involved in the breeding goal. The number of 10 to 1 5 traits may usuallyTable 4. Marginal

profitsof the traits in the breedinggoal

V (Y u a n )litter siz e b o rn a liv ed a ily g a infe e d c o n v e rsio n ratio k g /k gle an m e a t %wp Ha tev ra lu
h oeld in g c ap a city %

oa

V aA

9 4 .3 2
0 .4 9‑1 0 8 .0 0
2 .8 6
1 0‑05 .0.3 05

0 .3 6 8 7
3 2 .3 3 6
0 .2 4 4 92 .2 6 2 7
01 .7.0 06 89 90

3 4 .7 7 6
1 5 .8 4 5‑2 6 .4 4 9
6 .4 7 1‑07 .5.2 94 85

9 .1 3

0 .3 7 9 6

3 .4 6 6

in tram u sc u la r fat %Table 5. Performance of new dam lines of the Chinese

litte r siz e
ag e a t fin is h in g * d a yfee d c o n v e rsio n ra tio k g /k g
le a n m e a t %''Body weight

Table6.

at finishingmeat type pig

D m

D IV

D V

D V T

D v n

14 .0 81 7 8 .7
3 .14
5 8 .2 0

1 3 .1 5
1 7 8 .0
3 .0 16 1 .2 8

1 2 .5 0
1 7 6 .03 .14
6 0 .0 3

1 3 .0 6
1 6 2 .0
3 .3 05 9 .5 6

1 5 .6 7
1 7 8 .0
3 .1 95 4 .3 1of line Dili,

DIV and DV was 90kg, and that of line DVI and DVII was 85kg.

Summary of crossbreeding

parameters

for reproductive

traits

am

dm

m ean

1 .1 6.5 4

B ritish 1
trait

m ean

a

d

F re n c h 2

N o . te a ts

1 5 .8 5a g e a t p u b e rty d .
N o . b o rn a liv e
litte r b irth w t. k g‑
1 2 .2 4
1 4 .0 8

‑
ns
ns

‑
ns
1 .1 6‑
2 .0 6nsm e a n b irth w t. k g
su rv iv a l p re ‑w e a nl.s . a t w e a n in g
l.w t. at w e a n k g
A D G p o st w e a n g

1 .2 1.8 3 1
1 0 .107 7 .14 17

ns.0 6 1
ns
5 .8
ns

.14
.0 4 3ns
l l .260‑.2 3‑.0 7 6
.8 0
‑5 .6
ns

ns
ns
2 .2 02 0 .1A D G te st/g ro w e r g
F C R k g /k g

5 90
3 .0 3‑1 1 8.3 9

81?
‑.1 5 ns

‑114 o*.1 5

pen D F I kg
k illin g o u t k g /k g

1 .7 6.7 5 8

‑.2 2
ns

.1 5
‑.0 0 9e st. le a n %b a c k fa t m m
EM A
cm 2

3 1 .8 8
9 .9 72 8 .13 ‑7 .7 1 n s

ns
ns

a

d

am

dm

1 .8

ns
‑5 0
1 .4
1 .6

14 5
l l .1
1 2 .5

ns
‑1 .0

ns
.7

‑5 1
1 .8
ns

1 .0 8

ns

.0 8

‑.3 3

.0 5

1 0 .14 5 .8 3
38 3

ns
ns
‑3 2

ns
2 .8
33

1 .3 5ns
ns

1 .3
1 0 .1
ns

‑

63 1

‑116
.3

187

ns

29

ns

‑

‑.0 1 91 .3 6‑

4 3 .8
‑

‑7 .9
5 .9

2 .3 3
3 .2 2ns

means:estimated midparent value of Meishan and Large White, a:additive
direct, d: dominance, am: additive
maternal, dm: dominant maternal gene effects, l.s.: litter size, l.wt: litter weight, ADG: average daily gain,
FCR: feed conversion ratio, DFI: daily feed intake, EMA: eye muscle area(longissimus
dorsi).
1.Haley etal (1994),
Lee and Haley(1994),
Haley etal (1992),
Serra et al (1992)
2. Bidanel et al (1991),
Legault and Bidanel (1992),
Bidanel (1993)
3. weights recorded at 21 days

50

be enough to describe the whole performance of a population or of a breeding product(Fewson?1993a).
The pig is an economic animal of prolificacy and large litter size andhigh growth rate. Its economic value is expressed mainly by reproductive performanceof the sow and growth performance of offsprings.
Therefore, reproductive and carcass
quality, growth traits are important traits in pig breeding programs.
Chinese pigs possess commoncharacteristics,
such as high prolificacy,tenderand delicious meat, hardiness, but also some disadvantages, such as, low growth rate, low
feed efficiency and low lean content. Consequently the breeding objectives for Chinese
pigs are to improve growth rate, feed efficiency and lean meat content, and meat quality.
The goal traits and selection criteria of Chinese native pigs are shown in Table 3.
3. Calculationof economic weights of goal trait
The economic value of a trait is the marginal profit obtained, through a unitchange of the trait, considered above the population mean when all other traits are held
constant. Thus the economic weight of a trait in the breeding goal is defined as themarginal profit which one gets fromone difference between marginal return and marginal
costs (Fewson, 1 993b).
Using the establishedprofit equations and the investigatedcoefficients
andvalues under the conditions of production and market in Sichuan, China, marginal profit
for traits in the breeding goal of Rongchang pig is shown (Table 4) (Wang, 1995). The
genetic standard deviation oa shown in Table 4 is used to compare the economic
importance of traits for different units. Voa indicates selection importance for goal traits.
The results shown in Table 4 indicate that growing traits were the mostimportant traits in the breeding goal of Chinese native pig breeds. The litter size iseconomically importance and has a large influence on the economic return from pig
production. The carcass quality plays also an important part in economic return.
Use of Chinese Pigs
Chinese native pigs have high prolificacy,fine meat quality
and stressresistance, but also have low growth rate, thick skin and low lean meat, and it is moredifficultto adapt them to the requirements of modern commercial pig production.
Therefore, in the future Chinese pig genetic resources can be used:-to develop new breeds or lines

51-to make gains through heterosis.
1. Development of breeds or lines
Since the founding of the People's Republic of China, many breeds/lines
havebeen developed. Improved pig breeds have a larger body size, daily gain of about 600g
and lean meat of about 50%. They are playing a great role in the pig industry in China.
In order to effectively
use our pig breed genetic resources and meet therequirement of commercial production, dam lines which are prolific and have lean meatmust be developed and used in cross breeding programs. Through selection over the last
decade, 5 dam lines have been developed and seem to offer good prospects for use incommercial production (Zhao, 1 995). Table 5 shows their performance.
2. Commercial use of Chinese pigs
Many countries have paid attention to the prolificacy of Chinese native breeds,
especially Taihu pig. Since 1979 France, the UK, USA, and Japan have introduced theChinese Taihu pig (Jiaxing Black, Meishan, Fengjing) and used it successfully
to improve
sowproductivity.
The results indicated that Taihu pig seems to offer the best prospect
for use in commercial crosses. The heterosis between Meishan and Western crosses arehigher than that of crosses between Western breeds. Results from France and the UK
populations are broadly in agreement in the litter size of crossbreeding
parameters, which
are summarized in Table 6 (from Mercer and Hoste, 1994).
In the USA,Young( 1994) evaluated reproductive traits of gilts and sows that are
one-half or one-quarter Meishan, Fengjing, Minzhu, or Duroc. First-cross Chinese gilts
were superior to first-cross Duroc gilts for nearly all reproductive traits. Meishan andFengjing first-crosses had similar reproductive performance and were better than Minzhu
first-crosses. Differences in reproduction among backcross breed groups were smaller
than among first-cross breed groups. Most differences among backcrosses were notsignificant but were generally numerically better for backcross Meishan and Fengjing
than for backcross Duroc and Minzhu.
The strategy for exploiting Chinese breeds is to develop a synthetic population
using European animals which are both lean and prolific. This would minimize the loss
in prolificacy, enabling the production of an F l parent female containing 1/4 Chinese
genes, which is still prolific, and may have somedisadvantage in carcass quality. This

52

strategy has been used by NPD(National
Pig Development Co. , UK) in the developmentof a synthetic population (Upton Meishan).
A synthetic population between prolificwhite breeds and Meishan had a clear advantage in litter size of some 3-4 piglets relative
to the Large White and Landrace sows. Synthetic pigs grow slowly, are less efficient andmorethan 50% fatter than white breeds at present. Predictionsof performance for a 1/8
Meishan slaughter pig suggest that relative to other NPD stock, these pigs will grow justas quickly, and produce an extra 1.5-2mm backfat at UK slaughterweights(Mercer
andHosk,,,994).

Meishan can be used successfully commercially, with an economic advantage inmost markets. If the whole of the UK industry were to adopt this strategy, then thebenefit would be about $45 per sow(Mercer and Hoste, 1994).
The above results ignore other likely benefits from the use of Meishan crosses,
such as an improvement in general reproductive performance, and possible adaptability
to different farrowing environment (English et al. , 1990; Legault and Bidanel, 1992).
There is also evidence to suggest that the Meishan may have a fibre which can be usedin the diet more efficiently(Edwards et al, 1991).
It is also possiblethat rapid
improvements in the carcass can be made in synthetic
populations
relative toconventional white pigs (Navean et al , 1992).
Acknowled gemen t
I should

like to thank Prof.

ChangshinWu for reading

IP., Caritez, J.C. and Lagant,H., 1990.

Ovulation

the manuscriptand giving

suggestions.

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P.Q.,

1992.

Xiang

pig:

a superraw material

ChinesePig Breeds.

Harbin,pp. 525-529.

Mercer, J.T. and Hoste, S., 1994. Prospects
Genet. Appl.

Livestock

Prod., Guelph.

Navean, J., Ducos, A., Bidanel,

J.P. and Bazin, C, 1992. Results

growth rate in the Sino-European
Harbin,Tiameslan

Proc. Int. Symp. Chinese

on eight
lines.

generations

Harbin,StudiesPig Breeds.

on some properties

Harbin,of Chinese

pigand Lee-Sungminiatureexperimental

pig in Taiwan.

Proc. Int. Symp. ChineseF., 1992.
Pig Breeds.

Development

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Proc. Int. Symp. Chinese

Mechanisms
Harbin,ofBama Xiang pig. Proc. Int. Symp. Chinese
Wang, P.Q., 1995. Effects
Ph.D. Thesis,

and strategies

of long-term
University.

Wang, R.X., 1990. Reproductive

characteristicsof the Fengjing

The highlyof high

Preliminary

selection

Agricultural

Pig Breeds.

embryo survival
studies

Harbin,on breeding
pigs.

nucleus

Chinesein localPig Symp.,

Harbin,pp. 37-51.

of Chinese

NativePigs.

Zhejiang

Sci. Tech. Publishers,

and one-quarter

Meishan,Fengijng,

and sows. Proc. 5th World Congr. Genet. Appl. Livestock
Zhang, Z., 1991.

ChineseTaihu Pig. ShanghaiZang, Z.G., 1986.

Pig Breeds

in China.

Sci. Tech. Publishers,
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Zhang, Z.G., Huang, W.Y.,Luo, M. and Gan, M.H. 1990.

Practical

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MinzhuProd., Guelph,

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Shanghaiin Chinese

Toulouse,

Pig Breeds.
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Young, L.D., 1994. Reproduction

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pp. 539-549.

prolificZhao, S.G., 1995.

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Xu, Z.Y., 1989.

mini-pig.

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Wang, A.D., Lan, G.Q., Guo, Y.F., Qin, S.H. and Li, B., 1992.

Xu, S.Q., 1992.

for leanPig Breeds.

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M., Bazer, F.W. and Martinat-Botte,gilts.

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Henan

Characterization
and Conservation
Resources in Indiaof Silkworm Genetic

RAJAT KUMARDATTA
Central

Sericultural

Research and Training

Ministryof Textiles,

Government of India, Mysore-570008,

Institute,

CentralSilk Board,
India

Abstract
India has the distinctionsilk vis, mulberry,of being the only country to produce all commerciallytasar (both tropicalmulberry silkworm Theophila

and temperate),

eri and muga. Oak tasar Antheraea

sp. are among the wild forms seen in the Sub-Himalayan

variety Antheraea mylitta is found in as many as 1 8 eco-races in different
riciniis a domesticatedsilkworm

assama,a semi-domesticateddistributedin the North-Eastern

eco-climaticStatessilkworm (muga) produces the costliest

were initiatedin 1980s.

A Centralized(Tamil Nadu state) maintains& Training

Institute(Assam state)(Biharand morphologicalis maintaining

The characteristics

Germplasm Stationa large number of indigenousat Ranchipreserve their genetic

and conserve different

state)

silkworm

Similarly,

Centralof silkworm

breeds and races are limited
For mulberry

divergentusing

DNA level polymorphism,

useful biochemical
the approachand biometricfor characterizationin different parts of India
in the hybrid form.

silkworm

establishednichesin Hosurregional

Muga Research Station

niches toat Bokothe muga races.
various commercial traits.

electrophoresis,

Philosamia

whereas Antheraea

breeds in their eco-climaticfor mulberryof larvae and cocoons, besides
geneticallytasar

and exotic breeds, and Central Tasar Researchtocharacterize

and theTropicalgolden silk and is endemic to Assamconserves the tasar eco-races in different

characteristics.

ofroylei,belt.
regions.

of India,state. Many indigenous races of mulberry silkworm, Bombyx mori are still available
in addition to hundreds of newly evolved breeds that are commercially exploited
Steps to catalogue, characterize

known varieties

populations

DNAfinger printing,techniques

for characterizing

and conservation

to voltinism,molecular

morphologicalfeaturessilkworm breeds, attempts

are made

genetic

techniques,etc. The paper provides
the divergenceillustration

isozymefor theseof the silkworm breeds andof silkworm breeds, genotypes

and important

genes.

1.0. Introduction
Sericulture in India is as old as Indian culture. According to the Western history,
mulberry culture spread to India by about 140 B.C. from China. But this belief has beenchallenged
by many Indian scholars.
According to Mukherjee (1919),
the Aryans
discovered silkworm in the sub-Himalayas beyond Kashmir. There are many references
fromancient Hindu literature like the Rigveda, Ramayana, Mahabharatha on the use ofsilk clothes and sericulture (Nanavaty, 1990).
India has many species of sericigenous
insects and the flora of their food plants. India is the only country having the tradition

56

of using four silkworm varieties vis, mulberry silkworm(Bornbyx
rnori), tropical andtemperate tasar (Antheraea mylitta and Antheraea roylei), eri (Philosamia
ricini) andmuga {Antheraea assama). The ancestral silkworm variety, Theophila sp. was also
found in the Himalayan belt. Prior to the 1 9th century, sericulture was mostly confined
to Northern and Eastern India and only during a later period, it spread to the Southern
peninsula(Nanavaty, 1965).
Presently, 90% of the Indian silk is produced from thedomesticated mulberry silkworm Bombyx mori and only 10% of the silk comes from nonmulberry silkworms.
Despite the long history of sericulture in India from time immemorial, nothingis known about the characteristicsof the ancient races except that the reared silkworms
produced white and colored cocoons. Prior to the British period ( 1 8th century), silkworm
races were chiefly produced and maintained by the egg producers (Datta, 1984).
The
research work to characterize the races and to prepare manuals on mulberry cultivation,
cocoon production etc. started in the British period. It is only in independent India, thata concerted effort was initiated for characterizing
and conserving the silkworm races andtheir populations
in different agro-ecologicalniches when the Government of India
constituted the Central Silk Board under the Ministry of Commerce in 1949.
2.0. Non-mulberry Silkworms
The natural silks are broadly classified as mulberry and wild or non-mulberry.
The non-mulberry sericulture includes tropical and temperate tasar, eri and muga. India
produces over 1 000 MT ofnon-mulberry silks every year. Being the homeland of several
sericigenous insects, the Sub-Himalayan region of North-Eastern India is believed to besite of origin for various lepidopteranspecies including those of commercial silkworms
vis, eri, muga and temperate tasar. Several ethnic groups have been are engaged in
productionof different varieties
of silk for a long time. The availability
and theexploitationof sericigenous
fauna in India with brief information
about these
lepidopteranspecies is presented (Table 1 ).
2.1. Tropical Tasar Silkworm
Indian tasar silkworm, Antheraea mylitta D. is one of wild fauna of tropicalIndia. It exists in a wide range of geographicalconditions and about 1 8 ecological races

Table 1. Availability

and exploitationof sericigenous

fauna in India

are recorded in different parts of the tropical forest (Table 2). In order to assess thegenetic potential of tasar silkworm, many surveys were conducted by the Central Tasar
Research & Training Institute (CTR&TI),
Ranchi since 1964 in different tasar producing
states (Jolly et ah, 1968). The tasar zones are located between 16° (E) latitude and 60°88 ° (NS) longitude.
These ecological populations/races
are either univoltine, bivoltine
or trivoltine and the voltinism depends on climatic conditions of the region. It may also
be seen from Table 2 that the ecological races/populations
are confined to the various
food plants. For example; Raily, Laria, Modal and Nalia are sal (Shorea robusta)
based, while others are Terminalia based and a few others like Seoni and Tira are
Lagestromia based.

Table 2. Eco-races of trooicaltasar (Antheraea

mvlitta)and their characteristics

(MnaaTable 3. Comparative

analysisof traits

T raitsin temperate

A . roy le itasar silkworm species

A . p e rn y i

A . p r oy le i

A .y a m a m a i

E g g c o lo u r

B lu ish g re e n

B ro w n ish

B lu ish g re e n

P in k ish /B ro w n ish

L a rv a l c o lo u r(N e w ly h a tc h e d )B la c k

D u ll b la ck

B la c k

Y e llo w ish g re e n

M atu re d la rv a l w t.(g )12 .0

1 2 .01 5 .01 6 .0C o c o o n stru ctu re

D on uebr leS h e llIn

S ine lareg leb leS ine lareg leb leS ine lareg leb lere e la b leO u te r sh e ll
u n re e la b le
C o c o o n c o lo u r

In n e r‑c re am ,o u te r‑lig h t

L ig h t
b ro w n

L ig h t
b ro w n

G re e n ish
y e llo w

C o c o o n w t. (g )5 .3

5 .2 8

6 .2 2

6 .0 0P e d u n c le

T h in , s h o rtw ith rin g

T h in , s h o rtw ith rin g

W e a k , sh o rtw ith rin g

T h in , sh o rtw ith rin g

S h e ll w t. (g )0 .5 00 .6 00 .7 6

0 .8 0F ila m e n t le n g th (m )

468

4 11

750

600

D en ie r

5 .0

5 .0

5 .0

5 .5

Tahlp

4.

Seasonal

variationin vieidParam eterof A nxsnmn

silkworm^

M inim um

Fecundity
H atching %Totallifecycle (days)
Larval span (days)
C ocoon w eight(g)Shellw eight(g)Shellratio (% )Filam entlength (m )ER R (% )5Cnnmmercial
domestionted
Tsihlptraitstraits102 (sum m er)
19.0(‑do‑)
50 (‑do‑)22 ( ‑do‑)
4 .1 (W inter)0.28(‑do‑)5.49(‑do‑)204 (‑do‑)
20 (‑do‑)ofFood P lant
Fecundity
L arvalw eight
Larvae spotted and unspotted.
C ocoon w eight
Shellw eight
Shellratio

eri sikworm

M axim um
240 (A utum n)
84 .0(‑do‑)120 (W inter)60 (‑do‑)
5.8 (A utum n)
0.57(‑do‑)8.39(‑do‑)
500 (‑do‑)
60 (‑do‑)varietv

Polyphagous (C aster,T apioca,Papaya etc.)
300 to 400
8 to 10 g
T ubercles w ith setae allover body.
3 ‑4 g0.4 ‑0.5 g
13 ‑ 14 %60

2.1.1. Characteristics
The commercial characteristicsof the various eco-races, presented in Table 2,show that many commercial traits like cocoon weight, shell weight, shell percentage aswell as denier vary significantly
amongst different eco-races of tasar silkworm raised indifferent regions. High genetic coefficient of variation has been reported for these traits
by various authors (Sen et al, 1976 and Chatterjee
et al, 1983).
The characters likelarval duration, shell weight, cocoon weight and shell ratio show high heritability
withlow genetic advance possibly due to inter and intra-allelicinteractions(Siddiqui
et aL,
1985, 1988, 1989).
2.1.2.

Cytologicalcharacteristics
Chromosome numbers in A. mylitta D. was studied by Sinha and Jolly (1967)
and the haploid number of chromosomes (n=3 1) was established.
Interestingly,
novariation was recorded with respect to chromosome number amongst different eco-racesof tasar silkworm. However, variability
at the cytologicallevel was reported by Gaur(1986)and Sinha et al (1994)
in many different eco-races with respect to chiasma
frequency. Gaur (1986) has also reported variation on the basis of occurrence of the B
chromosome.
2.1.3. Biochemical
characterization
Variability
at the biochemicallevel was studied through gel electrophoresisforsomeof the major eco-races ofA. mylitta. Out of 26 bands reported, the band numbers
3, 6, 17, 20 and 24 were commonin all, while bands 23 and 25 were found to vary in
different populations(Sinha et al , 1 994).
2.2. Temperate Tasar Silkworm
An extensive survey of the sub-Himalayan belt in India conducted in 1966 by
the CTR&TI, Ranchi indicated the presence of vast wealth of oak in these regions as wellas the occurrence of the indigenous species, Antheraea roylei. Oak tasar plant is also
found in the North-Eastern region of India covering the States like Manipur, Assam,
Arunachal Pradesh, Meghalaya, Mizoram and Nagaland. CTR&TI, Ranchi successfullyraisedthe hybrid from A. roylei and its Chinese counterpart ^, pernyi. The resulting
hybrid called A. proylei Jolly, proved to excel both parents in all economic characters andthus gave rise to temperate tasar culture in India. This industry is spread mainly in61North-EasternStates owing to ideal socio-economic

conditions,speciallyin Manipur.

2.2.1.

Characteristics
The distinctive
morphologicalcharacteristics
of different
temperate tasar
silkworms are presented (Table 3). A. proylei is more similar to A. roylei with respectto egg color and larval body color. But many characters of A. pernyi like larval size,
cocoon weight, shell percentage, filament length are similar to that of A. proylei. The
filament length in A proylei was found to be longer than that of A. pernyi and only single
shell occurred instead of the double shell structure seen in A. roylei.
2.2.2.

Chromosome number
The haploid chromosome number of these Antheraea species of temperate zone
have been determined {A. pernyi n=49; A. proylei n=30; A. yamamai n=31; A. frithii
n=31; A. sivalica n=31 and A. polyphemus n=30). The chromosome number ofA.
roylei shows polymorphism
in Eastern India varying from 32 to 34 and wheninterspecifichybridization
was performed between A. roylei and A. pernyi, variable
chromosomenumberwas seen in the progeny. However, in the surviving A. proylei, thechromosome number was found to maintain n=49. Nagaraju and Jolly (1986)
have
shown how the chromosome orientation takes place in A. proylei.
2.3. Muga Silkworm
Muga silkworm is cultivatedonly in Assam because of its characteristic
ecological requirement of host plants. The polyphagous muga silkworm feeds primarilyon the leaves of Machilus bombycina (som) and Litsaea monopetala (soalu).
The
Eastern Goalpara and South-Western part of Kamrup District of lower Assam are themajor seed areas and the trade is largely in the hands of the tribal community.
Commercial rearing is practiced mainly in Sibsagar and Lakhimpur and to a lesser extent
in Nowgang, Darang and other Districts. The naturally grown cocoons are used for seed
and over 80% of the commercial cocoons produced in the plains are reeled on a large
scale in the Sualkuchi area.
2.3.1.

Characteristicsof Muga
The range of variation in commercial traits of muga silkworms reared in different
seasons is presented (Table 4). The egg number and the hatching percentage are lower

62

in summer compared to autumn. In winter, the total life cycle extends to 120 days
compared to 50 in the summer. Similarly, the cocoon weight, shell weight and shell ratio
are reduced very much in summermonths compared to those reared in winter. The larval
body of the muga silkworm is characterizedby black inter segmental markings over theyellowish
body. At maturity, the larval body weight reaches 15-16g.
The body iscovered by tubercles on dorsal and lateral (upper and lower) sides. The tubercle setae arepointed and blackish in color.
2.4. Eri Silkworm
Eri silkworm {Philosamiasp.) is the only one in the non-mulberry category which iscompletely domesticated and represented by two species in India today. The NorthEastern state ofAssam in India is considered to be the original home of eri silkworm andits advent in India is lost in antiquity.
However, the earliest reference documented in
1779 speaks oferi silk produced in Ghoraghat of undivided Bengal. Eri silk is popularly
knownas poor man's silk owing to its cheaper cost of production and lesser market priceof the spun silk produced out of its unreelable cocoons.
2.4.1.

Morphologicalcharacteristics
Philosamia(Attacus) cynthia is still found in wild state and performs poorly
under domesticated rearing. It is either bi or trivoltine and produces compact, light brown
cocoons. Philosamia ricini is the domesticatedvariety reared in many North- Easternstates including
Assam, Bihar and Orissa on the castor (Ricinis sp.) leaves.
But
ericulture is being practiced in many isolated patches spread over other parts of India andexploiting its polyphagous nature, eri silkworm is also reared, though on minor scale, on
Tapioca and Papaya. The cocoons are comparatively
loose and mostly white but
sometimes brick red. Commercial traits of this domesticatedvariety are given (Table 5).
3.0. Mulberry Silkworm
3.1. Mulberry Silkworm in Wild
Theophila religiosa, a wild species of silkworm is considered as a candidate forthe ancestral form of the domesticated mulberry silkworm (Tazima, 1964). Theophila
species are still found widely distributedin their natural habitats
in India as apolyphagous silkworm. A recent survey has recorded its occurrence on mulberry in subHimalayan regions, especially
Kalimpong (West Bengal), Sikkim and Manipur. It was

63also found prevalent on Artocarpus, Ficus bengalensis(Singh and Pavan Kumar, 1995).

and Ficus religiosa

food plants

3.2. Domesticated Mulberry Silkworm
The mulberry silkworm rearing has been practiced since ancient times in India.
According to historians,
some indigenous races have come to India through Siam
(Thailand).
Unfortunately, not much is known about the descent of indigenous races in
the literature.
India became home land for the silkworms which were brought from China
and then at a later stage spread to Northern and to some extent, Eastern India. However,the scenario of sericulture, is well documented and presents a picture of the diversity ofsilkworms that Indian people reared since 1 8th century. "Handbook of wild silk of India"
describes six species from the genus Bombyx prevalent at that time.
3.2.1.

Characteristics
of mulberry silkworm
The multivoltine mulberry silkworm, race Nyapau, is prevalent in India and wasreared both in Burma and Bengal in 1 8th Century. It was often referred to as Burmese
silkworm and produces multivoltine
white and yellow cocoons. The races namely
Madrasi, Chotopalu, Cheenapalu, Mysore, Boropalu, Lahemia and Kashmir that werereared in 1 8th century in Eastern, Southern and Northern India are presented (Table 6).
The characteristics
of most of these races are more or less the same, producing small
yellow cocoons of low quality whose larval period extends from 25 to 35 days. Only the
Boropalu race was found to be univoltine producing larger cocoons. The cocoons arealso mostly white with floss and are pointed at one end. The Kashmir race was also
univoltine producing golden yellow and white cocoons with high silk quality (Table 6).
In early 20th century, sericulture continued in Bengal. In independent India, thelocal indigenous multivoltine
race Nistari maintained its popularity
over the degenerate
univoltine Boropalu race which was reared only for a limited period during spring andautumn and the synthetic breed, Nistid, which was developed in 20th century by
hybridizing
Nistari with Italian breed.
Nistid was an improvement in terms of
productivity
and quality, but it was not popular in Bengal and later became extinct.
The characteristics
of different multivoltine
breeds including races like Nistari,
Sarupat, Moria and synthetic multivoltine
breeds are presented (Table 7). The cocoon
weight of the races ranges from 0.8 to 1.3g and SR % ranges from 11 to 15% indicating

64

Table 6. Mulberryv a rie tysilkworm varietiesin Pre-independent

India

Oanld
exdeistanpstlarececp eo rtf

C h a rac te ristic s

R e m a rk s

N y apau

B1 8uthrm cae,nBtuery
n g a l.

yMa en llo
du ltiv
Bw uormcltin
o ceoseeo;nws .h ite a n d

Aa slsoB orefem byrrex adrtora c a n en e sis

NcM aista
andaryran si//

B17e8n0g a‑l8 1silk w o rm .
Mwce olaecull
stic
oltiv
ou nnity
osdltinyearellonehd(7o wtb rito
w;lliafib
ea8 thnrebcroesyr;wo; dth
mith
s/y
riv
o thler);
insssgAp(Ha oslsopBu utto.larefecrnro)inrrees‑ stillid to

wm eilkllyu nw dheiter h;olarv
t w aeea thm eilkr; ym wo thh ites

Bp oenp uglaa lr in

wdspaithayns c2ino5nwdspainyictesoinur.ssum ma rk
m ine rga;nladrv3a5l
D e si/C h o top a lu1 8 th c e n tu ry .
NBvfirst
m(Oaeistaayrildnety
gtro
ebari
rel,A
th
apothnica
fadeInss
n lam
d ia .)

M u ltiv o ltin e (5 b re e d s p e r
wy2 e5olloar);rm
d awyssm
scb,oluacll;
isho o nlo;s;
la
o se
rv
M agolothsp
ldsaednu sk y ;A lso refe rre d toa s Bu tto(H.f onrtu). n a tu s

scC inh oeata
enpaat/p a lu /B1 8enthg ac l,Ae n tussa
ry .m

cMvoa corietieu ltivo nosltin.in fee ;rio
V re ry
to semarlie
allraA(Hslso
Bu tto. sreinnfe)erredn s is to

M y s o re

In tro d u c e d ins17u lta
8 5 nb iny T ip p u

M u ltiv o ltin e (In tro d u c e d fro m
C h in a )A lso re fe rred a s
B . m er id io n a lis .

LB eohroe mp aialu /K a rn a ta k a .A lso
C u d d a p p a h (A P )an d C o im b a to re (T N ).
MABob1 rig
e8ssa
en
ath
Italianingipmace).ul,narnntu(Md ryay.

NdLypU eay
oarv
onllo
inrm
ivsteawVoaldltin;llyspinMa atesta
wno;ereachL3r)itea0hflorg‑4ebse0nusy
rdtdc;saaoLoyncemsodso(1esntimg0su;‑me 12s;

NaA slso
oBw .retex
exfetintorrerc dt. toK a sh m ir

RKexepapshoorte
mrteirdd.lytoUws ilknh iv
iteqoultincali
o ctyeo;o; GnO softwlde nen
ithd yaverkery
llocwog loooourdrededB pueidec aetomm aeicbexin
a dtin c t

E u ro p e a ro u n d
18 55 .

o r b la c k w o rm s I m o u lt.

197 8 .

that these multivoltine
races have very low silk yield. The filament length ranges fromonly 384 to 527 m. The denier in races like Nistari, Sarupat, Moria is very low comparedto other breeds indicating that very thin yarn is preferred by the industry. The low qualityof these breeds is reflected in the neatness which is low and floss percentage which ishigh.

Research institutesof the Central

Silk Board collectedmany hybridsfromTable 7. Characteristics

multivoltine

mulberrysilkworm varieties

R ace

oSrig
tf
o f inrL arv a l

C o c oFo n

C̲ o .lo rl｣(N 0o 0.)0

C m(g )

S(e
wh g)
ell
t

R(%
S Oilk.o)(mF L )

T ra d itio n a l ra ce s
N ista ri

W .B e n g a lM

S p in d le

G O Y

829 5

0 .9 3 4

12 .4

13 .3

P u re M y so re

K arn a ta k aP

S p in d le

GR Y

837 1

0 .8 6 7

12 .3

14 .2

‑do ‑
‑do ‑

P

P e an u t
S p in d le

W h iteG .W .

89 15
7320

0 .84 1
1 .1 1 6

9 .6
16 .1

C N ich i
S a ru p at
M o riaof some indigenousof India
D en .e r

R(%
S a^
. wk)

R e e,

p ,o ss492

1 .2 85 .0 28 4 .3

78

7 .7

352

2 .0 4

8 .0 07 4 .475

18 .6

l l .4
14 .4

375
4 32

1 .4 4
2 .0 85 .0 95 .4 57 6 .48 7 .0

73
77

4 .1
9 .8

A ssa mP

S p in d le

G .W .

8035

1 .0 8 8

1 5 .6

14 .3

4 30

2 .1

5 .3 28 5 .076

9 .5

N istid (Y )W .B e n g a lP

S p in d le

G O Y

9 0 150 .9 3 9

12 .7

13 .7

372

2 .4 2

6 .1 88 5 .477

5 .7

N istid (W )

‑do ‑

P

S p in d le

W h ite8150

1 .0 1 1

13 .9

13 .6

2 84

2 .5 35 .0

7 9 .7

79

8 .2

7 .4

E a rly bre ed s d ev e lo p ed fo r tro p ica l reg io n s
W .B e n g a lM

O v alG O Y

84 2 4

1 .14 3

1 6 .91 4 .83 75

2 .0 5

6 .0 18 8 .976

A 4E

G ‑rac e‑do ‑M

O v al

G R Y

80 52

0 .9 9 5

1 5 .5

1 5 .644 5

2 .2 2

7 .0 37 1 .177

9 .0

A 14 D Y
T .W h ite‑do ‑T .N ad uMP

O v al
S p in d le

G O Y
W h ite82 35
82 8 3

1 .1 2 8
0 .9 7 5

1 6 .614 .1

14 .7
1 4 .5

3 59
4 38

2 .7 6
2 .0 5

6 .3 6
7 .9 78 5 .78 6 .2

78
80

5 .9
9 .8

H o sa M y so re

K a rn atak aPO v al

G R Y

863 1

1 .2 19

1 9 .1

1 5 .7

3 51

2 .8 2

6 .4 98 2 .473

9 .5

K o la r G o ld

‑do ‑

P

O v al

W h ite843 9

1 .3 0 7

2 0 .3

1 5 .552 7

2 .2 3

7 .4 17 9 .4

80

5 .4

M U l

‑do ‑

O va l

G R Y

747 1

1 .1 7 9

1 8 .4

1 5 .640 5

2 .2 2

6 .0 38 7 .067

8 .1

(hibernatine")

Table

8. Genetic

Table

9.

diversity

Biochemical
India.

in the mulberry

parameters

silkworm

eermolasm stocks

used for characterisationof mulberrysilkworm

available

in India.

germplasm

stocks in

67

different
countries in the early 20th century and developed them into pure breeds toincrease the genetic resources. The genetic diversity in present day mulberry germplasm
stocks is presented(Table8). It may be seen from the synthetic stocks that thevariabilityof these breeds is very high for fecundity, fifth age larval duration, yield bynumber and weight (yield / 10,000 larvae).
Consequently, the commercial traits likesingle cocoon weight, shell weight, filament length were also found to range from very
low to high.
3.2.2. Biochemical
characterization
During the breeding process, a large number of silkworm eggs have been isolated
and stabilized
whose maintenance and use in India warrants proper classification
andlisting.
Attempts have been made to classify these races on the basis of bio-chemicalparameters along with other traits used for characterizationof mulberry germplasm
stocks in India. Both qualitativeand quantitative parameters viz. , isozymes, activity ofdigestive
amylase, digestive
alkaline phosphatase,protease, invertase, haemolymph
amylase, haemolymph glucose-6 phosphate dehydrogenase,haemolymph trehalose
content were studied to understand the diversity of breeds. The minimum and maximum
values of various bio-chemical traits used for characterizationof the silkworm breeds isshown (Table 9). Both diapausing and some non-diapausing strains show negligible
digestive
amylase activity at pH range of 3-1 1, while most non-diapausing strains
registeredstrikingly
higheramylase activity at pH 3 to 9 (Abraham et aL, 1992).
Polyacrylamide
gel electrophoretic
analysis revealed the occurrence of only anodal
digestive and haemolymph amylases in the diapausing strains, whereas both cathodal andanodal isozymes were seen in the digestive fluid of the non-diapausing strain.
Mulberry silkworm germplasm stocks also showed genetic variability
in termsof haemolymph esterase isozyme (both a and b) patterns.
A total of 12 bands with 3
bands each in the four (A, B, C and D) zones were detected, but none of the strains
showed all the bands. Bl and B2 bands were present in all stocks whereas the highmobility band Al was observed in only one and A3 in two stocks, respectively.
A2 wasfound in all the hibernatingstrains and may be associated with diapause.
3.2.3.

Biometricalcharacterization
In Central Sericultural Research and Training Institute,

Mysore, attempts

were

68

also made to classify stocks by Mahalanobis genetic divergence techniques(Raoet al 91989;
Jolly et al, 1989). In addition, a detailed analysis was undertaken to test theefficacy of hierarchicalagglomerative clustering (UPGMA method) in grouping the races
and strains of the mulberry silkworm, Bombyx mori L. The analysis was based on datafrom 54 selected races/strainsof different
geographic
origins with varying yieldpotentials(Chatterjee
and Datta, 1 992). The results indicate that seven clusters can befound with yield parameters alone, whereas the inclusion of biochemical parameters inclustering resulted in only two broad groups : one having all the breeds with high cocoon
weight and shell weight and the other having all the low yielding silkworm strains.
Further sub-groupings under these two groups highlights
genetic differences associatedwith the differentiationof various groups of races in temperate and tropical races as wellas their significance for silkworm breeding (Chatterjee
and Datta, 1992). This study
represents a novel approach to analyze the relationshipof different silkworm races andbreeds on the basis of Squared Euclidean distance. It suggests the prospects of easy
handling of a large number of silkworm races and strains and supports the observation
onclustering system proposed by others (Peters and Martinelli,
1 989). The results also
highlighted
the use of biochemical parameters in obtaining more desirable information
on genetic relationships.
3.2.4. Characterization
using DNA markers
Techniques of studykig DNA polymorphism by RFLP, minisatellite
DNA and
RAPD markers have been recently initiated to further characterize our mulberry silkworm
germplasm stocks. DNA profile by RAPD technique of 1 3 silkworm genotypes of diverse
origin were studied by generating 2 16 amplified products using 40 random primers(Nagaraja
and Nagaraju, 1995).
Pair-wise comparison of amplified products andhierarchial clustering on the basis of Nei-Li Similarity Coefficient resulted in two distinctgroups of six diapausing and seven non-diapausing genotypes which suggests that RAPD
could be used for characterizationof silkworm stocks. Further, DNA-finger printing ofsilkworm strains by first digesting
the genomic DNAs with BstN l and Hinll restriction
enzymes followed by hybridization
with banded Krait mini satellite DNA, Bkm-2(8),minisatellite
probe revealed 9-3 1 discrete intense bands some of which were specific to
the genotype (Nagaraju et al , 1995). This once again highlights
the potentialsof these
methods for varietal identification.

69

3,3. Conservation
3.3.1.

Non-mulberry
At the initial stage of the establishmentof the Research Institute at Ranchi, all
the eco-races were brought to Ranchi and maintained in the establishedbush plantationsof Terminalia tomentosa, T. arjuna and Shorea robusta. Later, it was realised thatmanycommercial traits had deteriorated
or modified due to the maintenance of different
geographicalpopulations
in a new niche. Hence, the Central Silk Board decided thatthese eco-races are to be maintained in their respective ecological niches where they wereavailablefrom time immemorial. At present, CTR&TI, Ranchi is maintaining
these
breeds through different Basic Seed Multiplication& Training Centres (BSM&TC)
andRegional Tasar Research Stations establishedin different states of India. The cocoonsof different ecologicalraces are also preserved in the forest areas without seasonalcollection as practiced by the tribals.
It has been found that restricted multiplicationofthe eco-populations
in different pockets have strengthened the availabilityof A. mylittain the various regions of the country. The tribals, who are involved in tasar culture aretrained periodicallynot to interfere in natural populations that are maintained in theforest.
For conservation of the wild^A. roylei as well as the synthetic species, A. proylei
in their respective eco-geographicalenvironments, Central Silk Board has establishedthree Regional Tasar Research Stations, one at Imphal in the North-Eastern state ofManipur; another at Batote (Jammu & Kashmir state) in the Western sector and the third
one at Bhimtal in the Northern state of Uttar Pradesh.
The conservation of Muga silkworm has received attention, as this species isbeing endangered. The Assam State Sericulture
Department has earmarked "Muga
village Grazing Reserves" which are being allotted to the muga rearers with a nominal
10 % levy. This may on the one hand, save the habitat of muga silkworm from being
depleted or diverted for any other purpose and on the other hand, the muga silkworm
rearers are encouraged to keep up the traditional
avocation. Central Silk Board had
established a Regional Muga Research Station in 1 98 1 at Boko (Assam) which has now
been upgraded to a full-fledged
Research Institute.
It endeavors to solve technicalproblems related to muga sericulture and thereby aid its conservation.
Central Silk Board of India has also established
an Eri Research Station at

70

Mendipathar in Assam which is maintaining
a number of lines to fix up morphologicalsegregants
and some epigenetic
variations.
It is also actively concerned with thecharacterizationand conservation of the wild P. cynthia species in its own habitat.
3.3.2.

Mulberry
For conserving mulberry silkworms, Central Silk Board has establishedanexclusive Germplasm Maintenance Station at Hosur (Tamil Nadu state). Besides, thegermplasm stocks are maintained at different centres of the Central Silk Board as
indicated below:
CentreMulberry and Silkworm Germplasm Station,
CSRTI, Mysore, Karnataka
CSRTI, Berhampore, West Bengal
RSRS, Kalimpong, West Bengal
CSRTI, Pampore, Jammu &Kashmir
RSRS, Dehradun, Utter Pradesh
RSRS, Coonoor, Tamil Nadu

No. of mulberry
Hosur, Tamil Nadu

silkworm

stocks maintained
226
200
24
52
129
24
30

3.4. Conclusion
To conserve commercial live stock of silkworm and genetic material, India hasestablishedthe Mulberry and Germplasm Station at Hosur in Tamil Nadu state for themulberry silkworm Bombyx mori. The collection at this centre exceeds more than 100
varietal
stocks of commercial hybrids and nearly 200 genetic stocks.
The uniquedistinction of India for its rich fauna of sericigenous insects of non-mulberry crop is wellknown. North-Eastern India including states of Assam, Arunachal Pradesh and Manipur
has been identified as the natural habitat of nearly a score of sericigenous
insects. Manyof these fauna are threatened with extinction.
It is suggested, therefore, to have anInternational centre for collection and maintenance of silkworm live stocks, characteristicof this region. Possibilitiesof having regional centers in other Asian countries especiallyChina and Japan is proposed. A special emphasis should also be made on collection andpreservation of a number of non-mulberry silkworms especially,of the Anaphae genus
in Africa.

71
References
Abraham,E.G., Nagaraju,

Chatterjee,

J. and Datta,

R.K. 1992.

Biochemical

studiesanalysisin diapausing

and non-diapausing

Bombyx mori L.;

Comparative

Biochem. Mole. Biol.

22(8):867-873.

S.N., Bansal, A.K., Vatsauliya,
studiesin tropical

March 10-13:
Chatterjee,

P.K., Sengupta,

tasar A mylitta

Bongalore,of amylases

A.K. and Sengupta,

D. National

Seminarin the silkworm,
strains.

K. 1983.

Insect

The variability

on Silk Researchand Development,of 54 races and strainsof the mulberry

India.

S.N. and Datta, R.K. 1992.

Hierarchicalclusteringsilkworm,

Significanceof biochemical

Bombyx mori L.;

parameters.

Theor.

Appl.

Genet.

85:394-402.
Datta, R.K. 1984.
Gaur, J.P.

Improvement of silkworm

1986.

Cytology

Sericologia,

26(2):

Jolly, M.S., Chaturvedi
Jolly,

races {Bombyx mori L.) in India.

of supernumery

A survey of tasar crops in India.

M.S., Datta, R.K., Noamani, M.K.R., Iyengar,
and Rao, P.R.M. 1989.

Studies24(3):393-415.

Drury (Lepidoptera).

divergence

IndianJ. Seric. 7(l):56-57.

C.S., Basavaraj,
in mulberry

H.K., Kshama, G.

silkworm,

Bombyx mori

29:545-559.

N.G. 19 1 9. Handbook

of Sericulture.

Nagaraju,

J. and Jolly,Interspecific

M.S. 1986.

reassessment.

Nagaraju,

M.N.S., Nagaraj,on the genetic

Mukherjee,

Nagaraja,

mylitta

14 1 -146.

and Prasad, S.N. 1968.

L. Sericologia,

Sericologia,

chromosomes in Antheraea

Government Book Department,
hybridsof Antheraea

Calcutta,

India.

roylei and A. pernyi - a cytogenetic

Theor. Appl. Genet. 72(2):269-273.

G.M.and Nagaraju,

J. 1995.

arbitary

Electrophoresis

primers.

Genome fingerprintingof the silkworm,

Bombyx mori using random(in press).

J., Sharma, A., Sethuraman,

B.N., Rao, G.V. andsilkworm Bombyx mori using

bonded krait

Singh,

minor satellite

L.

1995.

DNA fingerprinting

DNA-derived

probe.

inElectrophoresis(in press).
Nanavaty,

M.M. 1965.

Silk

from grub to glamour.

Paramount Publishing

Nanavaty, M.M. 1990.

Silk

production,

processing

and marketing.

J.A.

HierarchicalPeters,J.P. and Martinelli,
germplasm

1989.

collection.

Theor. Appl.

Rao, G.S., Das, S.K., Das, M.K. and Nandhi,
silkwormSen, S.K., Sengupta,
path(Bombyxmori).

IndianA.K., Das, N.G. and Jolly,

co-efficient

analysisclusterHouse, Bombay, India.

Wiley Eastern

analysis

Ltd, India.

as a tool to manage variation

Genet. 78:42-48.
S. 1989.

Genetic

divergence

J. Agric.

Sci. 56:767-765.

M.S. 1976.

Studiesand discriminal

functionsamong bivoltine

on the genetic
in Antheraea

variabilities,
mylittaraces ofcorrelations,

D. IndianJ. Seric.

15(1):9-14.
Siddiqui,Siddiqui,

A.A., Debnath,

A.K. and Sengupta,

and their

traitscontributing

A.A., Sengupta,

K. 1985.

in A. mylittaof A. mylitta

Variability

J). Sericologia,

A.K., Ajit Kumar and Sengupta,

of some charactersin

D. Sericologia,

and correlation

studiesof shellweight25(l):45-50.

K. 1988.

Genotypic

28(1):1

87-192.

and phenotypicvariability

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Siddiqui,

A.A., Sengupta,
effects

Singh,

A.K., Das Mahapatra,of some quantitative

R. and Pavankumar,
Theophila

T. 1995.

sp. Proc. Curr.

SeptembertraitsD.P., Ajit Kumar and Goel, A.K. 1989.

Directon silk yield

Collection
Tech.

throughand indirectin A. mylitta

D. Sericologia,

29(2):21

1-214.

explorationof mulberrysilkworm

resources,

CSR&TI,

Mysore,Sem. on Mul. Breed. Genet & Mol. Biol.,

20-22.

Sinha,

S.S. and Jolly,

M.S. 1967.

Chromosome

number in tasar silkworm

Sinha,

A.K., Sinha, R.K., Goel, A.K., Sinha, B.R.R.Pd

Antheraea

mylitta

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Tazima, Y. 1964.

aspect of tropicalThe genetics

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of silkworm.

and Thangavelu,
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myllitta

A review on the breeding

D. Proc. Conf. & Genet. 4:7-1 6.

Chicken Genetic Resources in Japan and New Evaluation
Research
KEIJIRO NIRASAWA1 , HIDEAKI TAKAHASHI1 , TSUTOMU RJRUKAWA1 *,KAZUHIRO KIKUCHI2, JUNKO NOGUCHI2, YOSHIAKI IZAIKE2* and TAKAO OISHI2^Department
Nationalof Genetic Resources I and 2Genetic

Instituteof AgrobiologicalKannondai 2-1 -2^ Tsukuba, Ibaraki*Present address;

Department

National

Instituteof Animal Industry

Tsukuba

Norindanchi

Resources II,

Resources,
305, Japanof Animal Reproduction,

P.O. Box 5, Tsukuba, Ibaraki

305, Japan

Abstract
There are many chicken genetic resources in Japan.
consists

They consistof the native breeds which were selected before 1 868.

after 1 868.

The third group are the breeds derivedof three groups.

The first group

The second group are the breeds imported

from crosses between native and introduced

Most of the native breeds were developed for special plumage,

breeds.

crowing or cockfighting.

Seventeen

breeds have been designatednatural treasures of Japan. Most of them have low egg and meat productivity.
Recently some native breeds have been used to improve chicken meat quality.
Japanese native breeds aredifferent

from commercial breeds based on gene frequency.

collecting

semen from native breeds and preserving

microsatellite

DNA for use as genetic

In the MAFF Gene Bank Project,

it in a frozen state.

markers in evaluating

chicken

We have initiatedgenetic

research

we areon chicken

resources.

Introduction
Chicken eggs and meat in Japan and many other countries today are produced
froma few chicken crossbreeds. Hens for egg production are crossbreeds between somelines of White Leghorn or Rhode Island Red. Chickens for meat production are mainly
crossbreeds between White Plymouth Rock and White Cornish. These factory chickens
are very high yielding, but they require a carefully controlled environment.
In contrast, traditionalor heritage breeds may have survival traits no longer
found in industrial breeds, such as, climatic adaptation and resistance to diseases.
These
breeds are an important reservoir of genetic diversity.
In Japan, there are many chicken breeds including industrial breeds. However
there are no indigenous chickens in Japan in the true sense of the word. Chicken breeds
in Japan were introduced into Japan at various times. These breeds have broad genetic
diversity and are important as genetic resources.

74Japanese

Chicken Genetic Resources
Japanese chicken genetic resources can be divided into three groups according
to their origin and when they were introduced into Japan(Obataet al , 1994). The firstgroup consists of the breeds developed before the Meiji Restoration of 1 868. Between
1635 and 1854, Japan was closed to all countries except for China and Holland. In that
period many unique breeds were developed for special plumage or crowing in Japan.
These breeds are called "native breeds". The second group are the breeds introduced after
1868. Many chicken breeds have been imported from America and Europe for egg andmeat production and to improve Japanese native chickens. These breeds are called"imported breeds". The third group are the breeds derived from crosses between thenative and imported breeds. These chickens are called "crossbreeds".
Japanese

Chicken Breeds
In Japan all native breeds and some crossbreeds are called "Japanese chicken
breeds" (Mitsui and Koyama, 1979; Okada, 1991).
Most of them are specializedforplumage, crowing or game. There are more than 30 distinctive
breeds (Table 1).
Seventeen of them have been designated as natural treasures of Japan. The estimatedpopulation
size of each Japanese chicken breed is shown (Table 2). Most Japanesechicken breeds have low egg production and meat yield. These breeds are reared in small
populations
by a few people as a hobby. Many of such breeds are in danger ofdisappearing.
Japanese chicken breeds retain genes of old Japanese chickens and areimportant as genetic resources.
Descriptionof Japanese Chicken Breeds
Onaga-dori (Mitsui and Koyama, 1979; Okada, 1991) (Figure 1)
Onaga-dori is a native breed. It is the most famous Japanese breed because
males have very long tail feathers. This breed can be found in Kochi Prefecture, Shikoku.
The main tail feathers of males do not molt and can grow by 90 cm each year. The tailfeathers sometimes grow longer than 8 meters, and the record is 12 meters To grow sucha long tail feather, special care in feeding and management is needed and the technique
is difficult.
This breed is thought to have originated from a mutation of Shokoku or
Totenko breeds, in the 17th or 18th century.

Table1. Japanese chicken

breedsG ro u p

B ree dN a tiv e B re ed s

KJC hitocInuhga roabnikk ,kkeeoIw
ai,* sh*,S,aOhiw
te‑jidS ib
nh aaoag*ktto
aodUri,kri,
oukri**oTT ksu,o,KkHksei*
uahinwch
imaa id
i‑jid
Mca h‑ojid
iniyri*oaoori,kh,riik
kS,Soai*aT*tsudo, okUT m‑hazoraig
aum d‑jidraejidao ruri*c ho o*ari,,,ri,
bJKoidO *Kookeuri*
inyMreoa inksh,w oSoai*dhh‑hoa,ikri,
migTicooejUhte*idata,nbo oki‑rio* *C ro s sb re e d s

T o sa ‑k u k in , K u m a m o to , N ag o y a , Iz u m o ‑c o c h in , M ik a w a , M iy ajid o ri

d e sig n a te d a s N atu ra l N atio n a l T re a su re
T a b le 2 . E stim a te d n u m b er o f c h ic k e n s
N u m ber
> 50 0 0
20 00 ‑ 5 00 0
10 00 ‑ 2 00 0
50 0 ‑ 10 00
5 00 >

B ree dS h am o , C h a b o
H in a id o ri, S a tsu m a d o ri, N a g o y a , U k k o k e i
T o te n k o , M in o h ik ic h a b o , U z u ra c h a b o , T o m a ru , S h o k o k u . G ifu ‑jid o ri
K o ey o sh i, M in o h ik i, T o sa ‑iid o rL U tai‑c h a n , M ik a w a
JIz
K ito
uu mrok kko oas‑c, ohK ciwah wina ,ac,TShoaiydsaao‑kk‑hkuig
ok,ejinM id, ieKo ‑jidri,u m Koamri,u reoIwktooa,dteTo ‑jidri,su sToh ri,
imo k uaO ‑jidc nh ai‑jid
go ri
a doori,ri,MS hiyibajida ttoo ri,
ri

modifiedfrom Okada, 1 988

Tomaru (Mitsui and Koyama, 1979; Okada, 1991) (Figure 2)
Tomaruis found in Niigata Prefecture, Honshu. This breed numbers about 1000
individuals.
This breed is famous for the prolonged crowing ability of males. The cocks
can crowfor up to 18 seconds. Tomaru is a large breed and the adult male body weight
is about 3.5 kg. This breed is thought to be derived from chickens imported from China
in the 16th or 17th century.
In Japan, besides Tomaru, there are two other long-crowing breeds, Totenko andKoeyoshi.
In Okinawa prefecture there is the Utai-chan with a crowing pattern very
different from the long-crowing breeds.
Shamo (Mitsui and Koyama, 1979; Okada, 1991) (Figure 3)
Shamo is a typical game type chicken, and is found all over Japan. Shamo is
divided into six varieties according to body size or shape. This breed is thought to be
derive from a Malay-type chicken imported from Thailand in the 1 6th or 1 7th century.
Recently Shamo is used for crossing to improve chicken meat quality.
The Satsuma-dori is similar to Shamo and is found in Kagoshima and Miyazaki

76

Figure1. Onaga-dori. Photo by Mr.Shichiro
Koyama was reproduced
from
Nihonkei
Taikan(Mitsui
andKoyama, 1 979).

Figure2.

Tomaru. Photo by Mr.Shichiro
Koyamawas reproduced from Nihonkei Taikan(Mitsui and Koyama, 1979).

£' å »'.;- ^^.^fhri^i,^

Figure3. O-Shamo the biggest
variety of
Shamo. Photo by Mr.Shichiro
Koyama was reproduced
from
Nihonkei
Taikan(Mitsui
andKoyama, 1 979).

Figure- ^ft***w"1.
*'-i*a*ftM*>'*.ss^^fr-35®^

4. Katsura-chaboa typical
variety of
Chabo. Photo was reproducedbycourtesy of Mr. Mitsuyoshi
Tatematsu
from Nihonkei
Taikan (Mitsui
andKoyama, 1979).

77

Figure5. Nagoya. Photo was reproduced bycourtesy of Aichi-ken
Agricultural
Research Center.

Figure

6. Gifu-jidori is a typical variety of Jidori.
Photo by Mr.Shichiro
Koyama was
reproduced
from Nihonkei
Taikan(Mitsui and Koyama, 1979).

Prefectures, Kyushu. This breed was derived from crosses between Shamo and othernative breeds in the 17th or 18th century. In Akita Prefecture there is the Hinai-doriwhich has a similar origin to Satsuma-dori.
Chabo (Mitsui and Koyama, 1979; Okada, 1991) (Figure 4)
Chabo is a bantam chicken. Its adult body weight is about 700g for the male.
In Chabo there are 26 varieties with different plumage patterns. The features of Chabo
are there small body size, large single comb, erect tail feathers and short shanks. This
breed seems to derive from chicken imported from Indo-China in the 16th or 17th
century. This breed is a hobby breed and is found throughout Japan.
Nagoya (Okada, 1991) (Figure 5)
Nagoya is a popular breed and was derived from crosses among indigenous
chicken of the Nagoya area, such as Buff Cochin and other imported breeds. This breed
has good egg production. Crossbreeds between Nagoya and White Leghorn were widely

78

used as layers until 1950fs. Since then the number of the Nagoya chicken has decreased.
Recently this breed has been used for meat production.
A similar breed is Mikawa. It was derived from crosses between Cochin andother imported breeds. The number of Mikawa chicken has declined because its meat
quality is not as good as Nagoya chicken.
Jidori(Mitsui and Koyama, 1979; Okada, 1991) (Figure 6)
Jidorimeans an indigenous
chicken.
Jidori retains primitive
chickencharacteristics.
Until the 19th century there were many local varieties of Jidori
throughout Japan. Many varieties of Jidori were crossed with imported foreign breeds
in the late 19th and early 20th century. Consequently many Jidori varieties became
extinct.
Now, there are three varieties, Gifu-jidori,Mie-jidori
and Tosa-jidori.
These
have been designated natural treasures of Japan. Besides these varieties some othervarieties can be found in various districts,
but these are reared in such a small numberthat they face extinction.
Use of Japanese Chicken Breeds
Most of Japanese chicken breeds are not used for food. Recently some Japanesechicken breeds, particularly
Shamo,Satsuma, Hinai-dori and Nagoya have been used to
improve meat quality (Okada, 1991).
They have been crossed with White PlymouthRock or Rhode Island Red and are used to produce special regional delicacies (Fig 7).
The meat fromthese crosses is considered better than that of the crossbreeds with White
Plymouth Rock and White Cornish (Oishi, 1 992).
Present

Status of Japanese Chicken Genetic Resources
The number of breeds and lines of Japanese chicken genetic resources is
estimated to be more than 200 including breeding stocks, experimental stocks andJapanese native chickens.
Most of the Japanese chicken breeds are reared by hobbyists
and prefectural
agriculturalexperiment stations. Their populations
are small and usually number lessthan 100. Hobbyists rear these breeds in very small numbers of two to ten pairs.
Breeding stocks are reared by commercial breeding companies, some prefectural
agriculturalexperiment stations, and two national livestock breeding stations. Most of
the breeding stocks are lines of four representative
breeds; White Leghorn, Rhode Island

Figure7* Distributionof regional chicken delicacies.
These prefectures are using the crossbreeds
breeds and foreign breeds.

with Japanese chicken

80

Red, White Plymouth Rock and White Cornish. In national livestock breeding stations,
there are 14 White Leghorn lines, 3 Rhode Island Red lines, 9 White Plymouth Rock lines
and 5 White Cornish lines.
Experimental stocks include many lines, such as lines establishedby selection
experiments, developed for disease modelling, or having a specific gene or trait. These
are reared by university laboratories,
such as, Nagoya University, Hiroshima University
and Tokyo University of Agriculture.
Some national research institutes,
such as, theNational Institute of Animal Industry and National Institue of Animal Health, rear someof them also. Besides these organizations,
there are some experimental animalcompanies.
There are also a few chicken breeds in small populations exhibited at zoos, parks
and shrines.
In Japan in recent years, the number of prefectural agriculturalexperimentstations and hobbyists has gradually decreased.
Japanese Chicken Breeds in MAFF Gene Bank Project
According to some studies on blood type and protein polymorphism, Japanesechicken breeds are different
from commercial breeds based on gene frequency(Hashiguchi ^a/., 1981; Okada ^a/., 1988; Tanabe er a/., 1991).
Japanese breeds havebroad genetic diversity
and are useful genetic resources. The Japanese Ministry ofAgriculture, Forestry and Fisheries (MAFF) started the MAFF Gene Bank Project in
1 985. In this project, the National Institute of Agrobiological
Resources (NIAR) has
been collecting
semen of various Japanese chicken breeds and preserves samples in afrozen state. To date semen is fourteen breeds and five varieties have been collected.
NewEvaluation Research for Chicken Genetic Resources
Blood type, blood protein polymorphisms,
and morphologicalcharacteristics
have been used to evaluate chicken genetic resources. These methods are insufficientbecause the number of genes that are involved for these traits is limited.
Molecular techniques offer many newtools to analyse genetic polymorphisms,
for example, Restriction Fragment Length Polymorphisms (RFLP), Variable Number of
Tandem Repeats (VNTR) which involve repeated short DNA sequences (Hillel et al. 91992). Microsatellites
are a type of VNTR and are tandem repeats with between one and81four bases. Microsatellites
are useful genetic markers (Hillel et al, 1992; Cheng andCrittenden, 1 994). Microsatellites
are evenly distributed
throughout the chicken genome,
they are highly polymorphic
and can be assayed using the Polymerase Chain
Reaction(PCR)
methodology, and they are inherited as codominant alleles in a Mendelianmanner.
To evaluate genetic characteristics
of chicken genetic resources, wehave initiatedresearch on the chicken microsatellite.
Takahashi et al. ( 1995) developed an efficientmethod to clone chicken microsatellites.
Microsatellite
markers can be used to assess
genetic distance between breeds or between lines, and to assess inbreedinglevels inpopulations, and for evaluation of genetic variation within lines (Hillel et al, 1 992). In
the future, microsatellite
DNA markers will contribute to linkage analysis of quantitative
trait loci (Chen and Crittenden, 1994). Research on the chicken microsatellite
is expected
to provide a new field for evaluation of chicken genetic resources.
References
Cheng, H.H.and L.B. Crittenden.
Science
Hashiguchi,

Microsatellite

markers for genetic

mappingin the chicken.

and E. Hiraoka.

1981.

Sciencedetermined
52:7 1 3-729.

J., EA. Dunnington
Poultry

T. Nishida,by the blood

proteintypes offowls.

Japanese Journal

DNA markers in poultry

breeding

1992.

T and S. Koyama. 1979.

T., H. Takeda and T. Oishi.
1994. Japanese native livestock
breeds.
information.
No. 13, FAO and UNEP, Rome, Italy, pp. 13-24

Oishi,

T. 1992.

Conservationand genetic

analyses.

of the Japanese

Pettoraifu,Tokyo,

Japan. (Japanese).
Animalgenetic

native fowls and their use for the regionalin the conservation,

Instituteevaluationof Animal Industry.

resources

delicious

meat

and use of livestockandMinistryof Agriculture,

Forestry

breeds. Report of the Grant-in-aid

for Scientific

Research

pp. 1 8-50. (Japanese).

Distribution

Hiroshima
Fertilizer

Taikan.

Reviews on recent progress(No. 60304037)
on preservation
Okada, I. 1991.

Nihonkei

genetic resources. National

and Fisheries,
Okada, I. 1988.

ofZootechnical

Review 4: 169-1 86

Obata,

production.

Phylogenetic(Japanese).

and P.B. Siegel.

Science

H. Higashiuwatoko

Mitsui,poultry

Poultry

73:539-546

T., M. Tsuneyoshi,
relationship's

Hillel,1994.

of Japanese

to Okada, I. from Ministry
and estimationof native

University,

Catalogue

Higashi

Hiroshima,of the native poultry

Technology

Taiwan, pp. 3-35

chickenof Education, Science
livestockand poultry

and Culture of Japan. Study
genetic resources in Japan.

Japan, pp. 1 06-1 09. (Japanese).
of Southeast

Asia (FFTC

Center for the Asian and PacificRegion,Book series No.43).
Taipei,Republic

Food andof Chinaon82
Okada, L, A. Shinjo,

Y. Yamamoto, S. Kimura and E. Hiraoka.

native breeds of chickens.
Okada, I. from Ministry
estimation
Takahashi,of nativeof Education,
livestock

Scienceand poultry

and Culture
genetic

Y., T. Iida,

A. Shinjo

and S. Muramatsu.

toStudy on preservation

andHiroshima

University,

method to detectmicrosatellite

Symposium
1991.

on Japanese(No. 60304037)

in Japan.

cloningstudies

Researchof Japan.

resources

Program and Abstracts of Beltsville
Maryland, USA. pp. 32

H. Yoshino,

Phylogenetic

for Scientific

Higashi
Hiroshima, Japan, pp. 1 10-120. (Japanese).
H., K. Nirasawa and T. Furukawa. 1995. Efficient
markers in chicken.
Agriculture,
Beltsville,

Tanabe,1988.

Report of the Grant-in-aid

XX, U.S. Department

Studiesofon the phylogenetic

relationshipsof the Japanese native fowl breeds 5. The comparisons among native fowl breeds
in Japan and its adjacent areas and European and American fowl breeds. Japanese Poultry Science
28:266-277.

(Japanese).

Questions

and answers in Session

1

To Dr. Zhanchiv
Q. Are there distinct differences in lines of sheep developed for different practical goals
such as use of meat, skin, wool? (Bodo)
A. No, there are no big differences. (Zhanchiv)
Q. I heard about the reintroductionof animals which had died out in Mongolia? Is the
reproduction proceeding well or are there bad effects of acclimitization?
Are they
wild or domesticated?
(Bodo)
A. They are acclimitizedwell in two regions. Most of them are from zoos and aredomesticated. There are some from Russia which are half wild. (Zhanchiv)
Q. What is the importance of different sheep products in Mongolia? (Bodo)
A. Meat and wool (for carpets) are most important. The skin of the Karakul breed isimportant too. (Zhanchiv)
Q. Howsuccessful has the reintroduction of Prezwalski horses been in Mongolia? (Tan)A. The horses from the Ukraine and Switzerland have performed satisfactorily
in thewild in Mongolia. (Zhanchiv)
Questions to Dr. Nirasawa
Q. In Indonesia we have someproblems with cryo-preservation of chicken semen. Could
you tell us the method you use and have you tested the semen for quality after sometime in storage? (Astuti)
A. Weuse the Lake's method for cryo-preservation. The motility is excellant. (Nirasawa)
Q. How many semen samples are taken to represent each Japanese native breed? (LiNing)
A. For each breed the number ranges from 10 to 500. The number of birds from which
semen is collected ranges from one to forty. (Nirasawa)

TECHNICALREPORTS

Session2

Genetic classification

Chairpersons
A. Amano
R.K. Datta

Genetic Classificationof Asian Buffaloes Based on
Biochemical and Molecular Polymorphisms
SOON GUANTAN
Departmentof Biology,

Universiti

Pertanian

Malaysia

43400 UPM Serdang, Malaysia
Abstract
Electrophoretic

analysisof 39 allozyme

Swampbuffalo from Malaysia, Indonesia, Thailand,of the Murrah breed of River buffalo.

Nei's genetic

between somepairs of genetically

Swamp populations

recognized

in developed

breeds of livestockloci was conducted on 1 5 populations
the Philippines,
distance

Australia,
estimates

countries.

of9 phenotypicallyand two Murrah populations

Swamp populations

base fragment of the mitochondrial
differeddiagnosticallywith the Murrah. Similar

results

were obtained

as the Murrah. Among the mainland

Asian Swamp at two sites.

Southeast

Nucleotide

populations.

microsatellite

sequencing

ofa 303a sequence that

The Lankan had the same genotype

Asian Swamp populations

whereas only one of these was present in the insularwhen a subset

were typed for 2 1 nuclear

among the populations.

cytochrome b gene revealed that the Murrah possessedfrom the Southeast

frequencies

as those between

An UPGMAdendrogram of three phenotypicallycluster

However, there was greater differentiation

based upon the allelicwere of the same order of magnitude

SwampLankan buffalo populations
loci.

of phenotypically

Sri Lanka and two populations

Nucleotide

four genotypes
sequencing

were observed

ofa 1 58 base pairfragment of the mitochondrial
D-loop from 80 animals revealed the presence of 33 genotypes.
These
genotypes separate into 2 major clusters. One major cluster contained genotypes found in the Swamp only
and the other cluster contained

genotypes

in both the Swamp and the River types.
are geneticallythat were found in the Swamp, the River (including

Lankan)Hence, both our DNA and allozyme data indicatethat the Lankans

River and that there is sufficient

warrant crossing between them as an alternative
Swamp(2n=48)

genetic

variation

present among the Swamp populations

to the usual buffalo breedingpracticeof River (2n=50)

and
to
and

crosses.

Introduction
The Swamp buffalo Bubalus bubalis,is indigenous
to Southeast China,
Myanmar,Assam in India, Indochina, Thailand, Malaysia, Indonesia, the Philippines
andSri Lanka (Cockrill,1987).
It is much closer to the wild buffalo than are the buffaloesof India. They are different from the Indian breeds not only in appearance but also inbehavior and use. In Malaysia, there is no wild buffalo and the habitat of the domestic
buffalo is swamp or marshland. Hence, they had been named Swamp buffalo in contrast
to the imported Indian Murrah buffalo which is called River buffalo (MacGregor, 1 94 1)
since their usual habitat is the river valley. Swamp buffalo has traditionallybeen usedfor draught power in Southeast Asia and the River for milk and meat. However, theSwamp is being increasingly
used for meat as Southeast Asia develops and in the U. S.
A., Swamp buffalo meat is being promoted as a health food since its calorie, fat andcholesterol contents are lower than those of beef cattle. While the River buffalo has been
selected to form improved breeds with high milk yield and differing in horn form, theSwamphas not been divided into different types. It has retained the low milk yield andprimitive horn shape of the wild Ami buffalo found in Northern India and Indochina(Cockrill,1987).
Chromosomal studies showed that the morphologicallySwamp
indigenous buffalo of Sri Lanka (the Lankan buffalo) share the same diploid chromosome
number of 50 with the River buffalo while the Southeast Asian Swamp buffalo had 48
chromosomes (Bongso et al , 1977).
Amanoand his team have put in a monumental effort to study morphological,
blood group and biochemical variation in several Asian livestock species over a periodof morethan twenty years (Amano et al , 1 992 and references therein).
However, since
they were studying many species it is understandable
that their results for biochemical
polymorphism studies in buffalo were based on sampling a relatively small number ofanimals from each locality (ranging from 2 to 28) and from a modest number ofbiochemicalloci (ranging from 8 in the earlier reports to 25 in the later ones). They
found that the number of polymorphic loci for buffalo per country ranged from 4 to 7.
Their results were published in separate reports for the livestock species of each country.
However, in their report for Nepal (Amano et al, 1992) they included a UPGMA
dendrogram of genetic relationships
based on the genetic distances of Nei ( 1 972) among
22 Asian water buffalo populations. This dendrogram showed that the 9 Southeast Asian
Swamppopulations clustered together while the 2 Lankan, 3 Nepali and 3 Bangladeshi
native buffalo populations
formed another cluster together with a Murrah population ofRiver buffalo from Sri Lanka. The Philippine
and Indonesian River populations formeda third cluster together with one Swamp X River hybrid population each from Indonesia
and the Philippines
respectively.
Hence, it is likely that introgressionof Swamp genes
had occurred into the Indonesian (Amano et al , 198 1) and the Philippines
River populations.
Apart fromthe work of Amanoand his colleagues little information is available
about the population
genetics of Southeast Asian Swamp and Lankan buffaloes.
Consequently,
we initiated this study in 1988 to elucidate the population
structure of

89water buffaloes through the use of biochemical and molecular techniques. We nowhave
data on the same animals for nuclear electrophoretic
and microsatellite
loci and for
mitochondrial
DNA cytochrome b gene and D-loop control region. I shall now give asummary of the results that we currently have from our study of Swamp, River andLankan buffaloes from Southeast Asia, Sri Lanka and Australia.
Materials

and Methods
Blood samples were collected from 15 Swamp buffalo populations between
1988 and 199 1 in Malaysia (three localities:
Trengganu, Sabah and Sarawak), Indonesia(Bogor, Medan and Sulawesi),
the Philippines(Mindanao),
Thailand (Chengmai,
Haadyai, Surin and KamPaeng Seng), Sri Lanka (South, North-central and Kandy) andAustralia (Northern Territory)
and from two populations of the Murrah breed of Riverbuffalo (Universiti
Pertanian Malaysia, Serdang, Malaysia and Kotaliya in Sri Lanka).
The samples were collected, processed and separated into plasma, red cells and buffy
coat, transported in liquid nitrogen to Kuala Lumpur and stored at -70°C (Tan et al,1991) until used.
The plasma and red cells were used for analyzing nuclear biochemical markers
by cellulose acetate, polyacrylamide and starch gel electrophoresis(Tan et al , 1 99 1) andpolyacrylamide
gel isoelectric
focusing (Tan et al, 1993).
The BIOSYS-1 computer
package (Swofford and Selander, 1989) was used to calculate Nei's genetic distances(Nei, 1 972) based on the allelic frequencies and to generate an Unweighted Pair Group
Method with Arithmetic Averaging (UPGMA) dendrogram (Sneath and Sokal, 1973).
DNAextraction, polymerase chain reaction (PCR) and cycle sequencing for a 303 basefragment of the mitochondrial
DNA cytochrome b gene were done following auser-friendly system (Lau et al. , 1995). This PCR and cycle sequencing system was also
used to sequence a 260 base pair segment of the mitochondrial
D-loop region using
primers derived from the known sequence of bovine mtDNA. However, due to unequal
reading of the sequences at the terminii, only a common158 base sequence which was
obtainable
for all the 80 animals typed were compared and 33 D-loop genotypes wereobserved. The frequencies of these genotypes were used to calculate the Da genetic
distances of Nei et al. (1983) among the 1 1 populations(same populations
with microsatellite loci data) from which the animal werd obtained and these genetic distance values
were used to cluster the populations by the neighbor joining method. The 33 D-loop

90

Table

1. Sequencing of a 303 nucleotide
Buffaloes form 14 populations
L o c alitie s/B re ed s

fragment of the mDNAcytochrome b gene of Asian Watern12

S e q u 3e n c e T y p e 4

5

L a n k a n , D o u th S ri L a n k a
M u rra h , K o ta liy a , S .L a n k a (n = 5 )M u rra h , U P M , M a la y s ia (n = 4 )S w a m p , T e re n g g a n u , M a ly a sia
S w an p , H a a d y a i, T h a ila n d
S w an p , K P S , T h a ila n d
S w am p , S u rin , T h a ila n d
S w am p , C h e n g m a i, T h a ila n d
S w am p , S a ra w a k , M alay sia
S w am p , S a b ah , M a la y sia
S w a m p , S u la w e si, In d o n e sia
S w am p , J a w a , In d o n e sia
S w am p , M in d an a o , P h ilip p in e s
S w a m p , N .T e rri to ry , A u stra lia

genotypes observed were also individuallyclustered using the proportion of nucleotide
differences and neighbor joining with the corresponding bovine sequence as the outgroup.
Twenty one microsatellite
loci were typed for using the method of Moore et al (1995)
from eight Southeast Asian Swamp populations namely Surin, Trengganu, Sabah,
Sarawak, Bogor, Sulawesi, Musuan and Australia; fromthe South Lankan population andfrom the Murah populations of Sri Lanka and Malaysia. The microsatellite
data wereanalysed in a manner similar to that used for the allozyme data.
Results
Data are presently available for 39 biochemical genetic loci in 1 7 populationsof which two are Murrah populations(Tan et al9 1994).
In addition, the threephenotypically
Swamp Lankan populations were shown electrophoreticallyto be of River
type since they clustered with the two Murrah populations on the UPGMA dendrogram
presented in Figure 1 (Barker et aL, 1991).
Hence, there are 12 geneticallySwamppopulations,
three from Indonesia, one from the Philippines,
four from Thailand, three
from Malaysia and one from Australia. In most populations,about 50 animals weretyped for the polymorphic loci and 25 for the monomorphic ones. Of the 39 loci, 14 werepolymorphic in Swamp populations and one diaphorase- 1 zone 2 was polymorphic only

91

in Murrah. Of the 14 polymorphic Swamp loci, glyoxalase- 1 was polymorphic in four
populations and the other 13 (acid phosphatase, albumin, amylase, catalase, diaphorase- 1
zone 3, group-specificcomponent, glutamate pyruvate transaminase,malatedehydrogenase, malic enzyme, peptidase-B, peptidase-C,
protease inhibitor
and transferrin) were polymorphic in at least nine of the populations.
The percentage ofpolymorphic loci ranged from 12.8 in the Northern Territory feral population to 35.9 in
Trengganu. Meanheterozygosity was also lowest in the Northern Territory population,suggesting that this population probably underwent a genetic bottleneck on introduction
to Australia.
Of the Southeast Asian populations, those in Sabah and Sarawak showed
lower genetic variability
than others. Measures of genetic variability
in the Murrah and
Lankan populations were similar to those for the Swamp populations.
The matrix ofgenetic distances among each pair of Swamp populations showed values ranging from0.002 to 0.058. Within countries, the highest value was for Malaysia (0.026) primarily
due to differentiationof the Sabah population. Our estimates of genetic distances among
Swamp buffalo populations were of the same order of magnitude as distances among
recognized livestock breeds in developed countries for example those for European
breeds of cattle (Gonzalez et al , 1987). The three Lankan populations were most closely
related to Murrah from Sri Lanka (D=0.011)
and then to Murrah from Malaysia(D=0.028).
This group of populations
was distinct(D=0.052)
from the Swamp
populations(Barker et aL, 1992).
About 20 animals from each of the 8 Swamp populations,
the South Lankan
population and the 2 Murrah populations were analysed for 2 1 nuclear microsatellite
loci(Barker ^ a/., 1995). All 2 1 loci were polymorphic with the number of alleles detected
per locus ranging from2 to 14 in Swamp buffalo and from 2 to 18 in River buffalo, with
one or more alleles at each locus shared between the two buffalo types. The mean
number of alleles per locus ranged from 2.6 (Sarawak) to 5.3 (Lankan buffalo).
Themeanobserved heterozygosity amongthe Swamp populations ranged from 0.5 89 (Surin)
to 0.400 (Sabah) while among the River group the Sri Lanka Murrah had the highest
value of 0.6 13 while the values for the Lankan buffalo and the Malaysian Murrah were
the same (0.53 1). The Australian population had a mean heterozygosity
value of 0.409
and 3.0 alleles per locus. Nei's (1972)
genetic distances (D) among the Swamp
populations ranged from 0.026 between the Surin and Trengganu populations to 0.492
between the Australian and Sarawak populations.
Among the River group the lowest D

92value (0.041)
was between the two Murrah populations
while the highest D value of
0.067 was between the Lankan and the Murrah population from Sri Lanka. The highest
D value among the 1 1 Asian buffalo populations
was between the Sabah Swamp
populationand the Murrah populationfrom Malaysia (1.125).
In the UPGMA
dendrogram based on the 21 microsatellite
loci and Nei's genetic distances (D) the
Lankan and the 2 Murrah populations form one cluster while the 8 Swamp populations
form another cluster (Figure 2).
Whengenetic distances (D) based on protein data for 25 polymorphic loci were
calculated for the same 1 1 populations with microsatellite
data available, it was observedthat the mean D amongthe 8 genetically
Swamp populations was 0.098, among the 3
geneticallyRiver (which included the Lankan) populations
it was 0.046 and for the
genetically
Swamp versus River populations
it was 0. 173. In comparison, the values
based on the 2 1 microsatellite
loci were 0.228, 0.056 and 0.897 respectively(Barker etal.9 1995).
Thus, microsatellite
loci showed much greater discrimination
among
populations in terms of larger genetic distance values than protein loci. MacHugh et al(1994) typed 1 1 microsatellite
loci in 6 European breeds of dairy and beef cattle namely
Aberdeen Angus, Here ford, Simmental, Charolais, Jersey and Friesian. The average ofall pairwise genetic distances (D) among these 6 breeds was 0.083. Therefore, themagnitude of genetic differentiation(D=0.228) based on 2 1 microsatellite
loci among the
8 genetically
Swamp Southeast Asian buffalo populations
is much greater than thatamongwell recognized and long-establishedcattle breeds (Barker et al , 1 995).
Our sequencing of a 303 base fragment of the mitochondrial
cytochrome b geneof Swamp buffalo from the three populations
from Malaysia, four populations fromThailand,two from Indonesia (Bogor in Java island and Sulawesi),
one from thePhilippines(Musuan in Mindanao island) and one from Australia and the two Murrah
populations
revealed that the Swamp and the Murrah River buffaloes
differeddiagnosticallyat two sites. The South Lankan population possessed the River genotype(Figure 3). Among the Southeast Asian Swamp four genotypes were observed in themainland populations
whereas only the most frequent one of these was present in theinsular populations (Sabah, Sarawak, Mindanao, Bogor and Sulawesi). All the nucleotide
substitutions
observed were transitions and silent.
Comparison of the sequences obtained from 80 animals for a 1 5 8 base fragmentof the mitochondrial D-loop region revealed the presence of 3 3 genotypes. Clustering ofFigure1.

Dendrogram of genetic distances among 12 swamp and 5 river buffalo
dendrogram is base on 39 allozyme loci (from Barker et al.^1991).

Figure2.

Genetic relationships
among 1 1 Asian water buffalo populations
mirosatellite
loci. UPGWA tree (from Barker et al9 1995).

populations.

This

based on 21 nuclear DNA

Figure3.

Phylogenyof 33 buffalo mtDNA D-loop haplotypes(BOV=bovine sequence as outgroup),using proportion
ofrnucleotide
differences
and neighbor joining.
*haplotype
found in river
buffalo only, **haplotype found in both river and swamp, and all others found in swamp only
(from Barker et ai, 1995).

Figure4. Phylogeny
populationfor 1 1 buffalo populations,(from Barker et al., 1995)

using mtDNA D-loop haplotype

frequencies

in eachthese 33 genotypes by the neighboring joining method (Saitou and Nei, 1987) resulted
in two major clusters (Figure 3). One major cluster contained genotypes found in the
Swamp only and the other cluster contained genotypes that were found in the Swamp,
the River (including the Lankan) and in both the Swamp and River types. When the
populations
were clustered based on the Da genetic distances (Figure 4) there weresimilarities
to the microsatellite(Figure 2) and allozyme (Figure 1) based clusterings.
However, the Swamp population from Bogor, Indonesia clustered with the three River(including Lankan) populations. But with 33 genotypes and sequences available for only
5 to 10 animals per population(average 7.3) the data are inadequate for reliableconclusions to be made. Hence, more animals per population and more populations
should be sequenced for the D-loop since our data showed the presence of a substantial
amountof variation and seemed to indicate that the Swamp is the ancestral type (Barker
etal,1995).

Discussion
Both our biochemical and molecular results revealed that the phenotypically
Swamp Lankan buffalo is actually of River type genetically.
The geographically96

separated Swamp buffalo populations of Southeast Asia are generally phenotypicallysimilar but there is sufficient genetic variation present among them for us to suggest to
buffalo breeders to consider crossing between animals from genetically distant Swamp
populations as an alternative to relying on crosses between Swamp and River buffaloes
in order to obtain hybrid vigour (Tan et al., 1994).
Moreover, crossing among the
Swamp would avoid any future fertility problem that may arise at the village level as aresult of the current widespread practice in countries like the Philippines(Steane et al ,1 994) of crossing between the River and the Swamp which differ in their chromosome
numbers. Our results emphasize the importance of conserving Swamp buffalo
populations
from different geographicalareas of Asia in order to preserve and use thegenetic variations present in them so as to improve the Swamp buffalo herds at thevillage and farm levels. In addition, our data support the hypothesis of Amano et al( 1 992) that the Swamp and the River buffaloes have been domesticated from different
origins and that mitochondrial DNA polymorphisms could be used to distinguishbetweenthem (Amano et al, 1994, Tanaka et al, 1995).
It is proposed that large scale population genetic studies such as ours and thoseof Amanoand his colleagues be extended to the Swamp populations of China, Myanmar,
Vietnam, Laos and Cambodia and to the River populations of India and Pakistan in order
to ascertain the levels of genetic variation present in them and to determine their genetic
relationships
to one another as well as to the buffalo populations which we have studied.
Also it would be desirable if the centres of origin of the Swamp and River buffaloes ofAsia could be identified.
Animals from more populations in mainland Southeast Asia
should be sequenced for cytochrome b and more populations as well as more animals per
population should be sequenced for the D-loop region. At the Department of Biology,
Universiti
Pertanian Malaysia, Serdang, Malaysia, we already have stored in our deep
freezers buffy coat samples from animals (random animals as well as those from complete families) and populations which ought to be, but which have not yet been, typed forDNAmarkers like microsatellites
and mitochondrial D-loop and cytochrome b. Although
they have all been typed using allozymes, we have subsequently demonstrated that DNA
markers are far more informative. Our current funding to do such DNA studies havebeen exhausted after weanalysed a subset of these animals and populations for the DNA
markers. However, it should be emphasized that overall the genetic relationships
obtained among the populations
that we studied are similar irrespectiveof whether

97

biochemical(protein-level)Thissimilarity

overallmeasuredor molecular

relationships(DNA-level)

markers

are usedgreatly

strengthens

theconfidence

that

among

the Asianwater

buffalopopulations

to obtain

can be placed
which

them.
on thewe havestudied.
Acknowledgement
I thank the Australian
PN 8364 and a project

Programme for Intensification
Ministry of Science, Technology
grant 50205-95-50.

Centre for International

grant Molecular

genetic

Agricultural
analysisof Research in Prority Areas (IRPA) throughand the Environment, Malaysia and Universiti

This report is mainly based on work done in collaborationof New England, Australia),

T.K. Mukherjee

and K. Yusoff (Universiti

Pertanian

Malaysia),

University

Australia).

The invaluableof Queensland,

Department staff from Australia,
collection

is gratefully

Research for fundingon Swamp buffalo

Indonesia,

and O.S. Selvaraj(University

the Philippines,

the Malaysian

grant 1-07-05-057
from thePertanian Malaysia throughwith J.S.F. Barker (Universityof Malaya, Malaysia),

S.S. Moore, R. Drinkwater
assistancethis work throughpopulations,and D.J.S. Hetzel (CSIRO,

of numerous University

Sri Lanka, Thailand

C.H. Lau

and Malaysia

and Veterinary
during sample

acknowledged.

References
Amano, T., Nozawa, K., Kawamoto,Y., Namikawa, T., Martojo, H., Abdulgani,

I.K., Nadjib,

H. 1981.

Morphological and genetical survey of water buffaloes in Indonesia. In: The origin and phytogeny
of Indonesian Native Livestock. Part II. The Research Group of Overseas Scientific
Survey.
Tokyo, pp. 31-54.
Amano,T., Namikawa, T., Maeda, Y., Tsunoda, K., Yamamoto, Y., Shotake, T., Nishida, T., Rajbhandary,
H.B. 1992. Constitutionof genes controlling
blood protein types ofNepalese water buffalo andtheir phylogeneticclassfication.
Report of the Society for Researches on Native Livestock 14:
89-100. Nagoya Foundation for Animal Science, Japan.
Amano,T., Myakoshi, Y, Takuda, T., Kikkawa, Y., Suzuki, H. 1994. Genetic variants ofribosomalDNA
and mitochondrial
DNA between Swamp and River buffaloes. Animal Genetics 25 : Supplement
1,29-36.
Barker, J.S.F., Mukherjee, T,K., Hilmi, A, Tan, S.G. 1991. Genetic identificationof strains and genotypesof buffaloes and goats in Southeast Asia (PN8364).
Final project report to the Australian Centre
for International
Agricultural
Research, Canberra.
Barker, J.S.F., Tan, S.G., Selvaraj, O., Mukherjee, T.K. 1992. Biochemical
genetic variation and genetic
distances among Southeast Asian Swamppopulations. In: Recent Advances in Animal Production.
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Phylogenetic
Relationships
Bubaline Species in Asia

and Conservationof Bovine and

TAKAO NAMIKAWA,KAZUAKI TANAKAand CHESTER D. SOLIS
School of Agricultural

Sciences,

Nagoya University

Furo-cho, Chikusa-ku,

Nagoya 464-0 1 , Japan

Abstract
A variety of wild bovine and bubaline
Asia.

These

speciesphylogenetically.
threatened

speciesare more or less related

However, most of these species

with extinction.

Detailedbiologicalstill inhabit

restricted

to the ancestor

areas of South and Southeastof domestic

cattleor water buffaloes

are thought

to be declining

in number and endangeredcharacterization

is inadequate

and schemes for conservationorare insufficient.
The wild species
subspp.),of bovine in Asia consistyak (Bos mutus) and kouprey (or Indo-Chineseof, the banteng(Bos javanicus),forest ox) (Bos sauveli).

gaur (Bos gaurus

The kouprey is now veryrare and little is known of its present status. No concrete genetic or reproductive data on the kouprey isavailable to compare with that of the related species, although very detailed morphologicalor osteometricalcharacteristics

have been described (Coolidge,

1 940).

phylogenetic

relationshipcattle,these wild speciesof domesticatedand domesticatedbubalis\(Anoa)

quarlesi)

Mindoro Island
In thissummarizing

the lowland

are generally

acceptedin the genus Bubalus;anoa (Bubalus(Figurepaper,

we elucidate

cattle(mostly1).

endemic to Sulawesi,

basic data are significant
to domesticatedoffspring(Figureof Philippines
our previous

because hybridof bovine can help elucidate

cattle are fertile

At least four wild species(Bubalus

The other wild species(Anoa) depressicornis)

Indonesia,

in the female)thebetweenthe Asian wild buffalo

and the mountain anoa (Bubalus

and the tamaraw (Bubalus

mindorensis)

endemic toand bubaline

species2).
phylogenetic

genetic

or molecular

in relation

to conservation

relationships

evolutionary

studiesof bovineof them. We also discussand use of these wild species,byhow these

which are closely

relatedor water buffaloes.

Phylogenetic
study on bovine species
Molecular evolutionary relationshipof bovine hemoglobin-ft varian ts
In native cattle populationsof Southeast Asia, three hemoglobin-P
globinvariants are generally found, namely HbP-A, Hbp-B and HbP-XBah. These variants arewell known to be expressed by the allelic genes, HbbA, HbbB and HbbX, respectively(Namikawa and Widodo, 1978).
We determined the amino acid sequence of HbP-XBah of the Bali cattle (Bosjavanicus) and compared it with two commonvariants, HbP-A and HbP-B, previously
reported in domesticatedcattle (Namikawa et al.,1983;
Namikawa et al.,1987).
The

B

100

lysinylresidue at Hbp-A1 8 was substitutedby the histidinyl

residueat the Hbp-XBali 1 8.

This change requires two DNA base substitutions
at the triplet codon. The HbP-A differsfrom HbP-B in the residues of 15, 18 and 119. This finding indicates that a common
ancestor of Hbp-B of the domesticated cattle and HbP-XBah of Bali cattle first diverged
fromHbP-A of the cattle through the Lys-^His substitution
at the residue 1 8. Bali cattle
has generally been accepted to be a domesticated form of the banteng (Bosjavanicus).
The evolutionary relationshipof HbP-A, Hbp-B and Hbp-XBah globin sequences
involves different allelic frequencies of the HbbA, HbbB and HbbX explainsdifferences between native cattle populations
in Southeast Asia.

genetic

Distributionof hemoglobin
globin alleles in native cattle populations in Asia
The geographicaldistributionof Hbb alleles in the cattle of Eastern Asia has
been described (Namikawa, 198 1)(Figure3). All of the native cattle populations
inSoutheast Asia are polymorphic with the HbbA, HbbB and HbbX alleles. The humplessor northern type cattle are nearly fixed with the HbbA, and the Indian zebu cattle was
highly polymorphic with the HbbA(0.64)
and HbbB(0.34).
The HbbX frequency was
uniquely high in the Bali cattle of Indonesia(0.79)
and decreased in populations distantfrom the Bali cattle population.
On the assumption of the three-way hybridization
process in development of the
Asian cattle, amount of gene-flow from Indian zebu (Pind), the Bali (PBal) and thenorthern type (PNor) were estimated for native cattle populationsof eastern Asia. P^orvalues dominated in the Japanese, Korean cattle and some Thai native cattle, while the
distributionof Pind, PBal and PNor in Southeast Asian native cattle suggests that the
Indian zebu cattle as well as the Bali cattle and northern type cattle have contributed tothese native populations
at various levels (Figure 4).
Genetic relationshipof cattle breeds/populations
using multi-genetic characters
Using genetic data on the nine blood group systems and three biochemical
polymorphic
loci, principal component analysis was conducted for many Asian cattle
populations(Namikawa et al., 1984)(Figure5). The results showed a large genetic
divergence between the northern type cattle (Korean, Japanese cattle and European
breeds) and southern type cattle (Indian zebu, many Southeast Asian native populations
and the Bali cattle). The southern type cattle are subdivided
into four major groups;

101( l )the Bali cattle, (2)Indian zebu, (3)Taiwan-Philippineand (4)Thai-Malaysia-Indonesia.
These genetic relationships
correlates to the Hbb allele distribution
mentioned above,that is, the Japanese- Koran native group is characterizedwith large P^r values, theTaiwan-Philippine
group with large Pina- PBal values and small P^or values, andThai-Malaysia-Indonesia
group with relatively
large PNor values and various Pina and
PBal values.
Source of genetic diversity in the native cattle populations of Southeast Asia andconservation
The above mentioned studies show the broad genetic differentiationof the native
cattle populations of South and Southeast Asia. It can be said that the broad wide genetic
variabilityof cattle is primarily provided by three distinct types of cattle or different"species" of bovine; probably Bos primigenius (the northern type or the humpless cattle),
Bosjavanicus (the banteng or the domesticatedform Bali cattle) and the zebu cattle ofwhich detailed ancestry remain unknown, Therefore, most of the genetic diversity of the
Asian cattle will be conserved in these three representative
types of bovine.
Phylogenetic
study on bubaline species
Phylogeny of Bubalus spp. by cytochrome b gene evolution
The whole of the cytochrome b gene region of 1 140 bp was sequenced for sevenbubaline mtDNAs; Synceros caffer (African buffalo), Bubalus mindorensis, Bubalus(Anoa) depressicornis, Bubalus (Anoa) quarlesi, and Bubalus bubalis (Sri Lankan wild
buffalo, river type buffalo and swamp type buffalo).
The pairwise sequence difference,
considering position of codons as well as transition and transversion were compiled.
These sequence data will be appeared in the databank of DDBJ and elsewhere. Only fivenucleotide substitutions(all transitionat the third position of the codon) were foundbetween the Sri Lankan wild buffalo and the river type buffalo. This substitution
rate iscomparable to that estimated within the river type buffaloes (Tanaka et al , 1995). On
the other hand, 28 substitutions
existed between the river and swamp type buffaloes.
The phylogenetic
tree of six Bubalus sequences was constructed using theneighbor-joining
method, in which Synceros and/or Bosjavanicus sequences were placed
as an outgroup. In the dendrogram (Figure 6), the swamp type buffalo and tamaraw madea cluster, but the river type buffalo and Sri Lankan wild buffalo were separte from thisFigure1. Genus Bos. Upper left: Gaur. Indian gaur ox (Bos gaurus gaurus) in wild (photyo by Dr. T.
Shotake). Upper right: Yak (Bos mutus). A domestic yak bull of Nepal (photo by Dr. T. Shotake).
Lower left: Banteng (Bosjavanicus).
The banteng bull at Surabaya zoo of Indonesia.
Thebanteng is a wild form of the Bali cattle. Lower right: Mishima-ushi(bull).
There are two purely
native populationsof cattle in Japan; one is Mishisma-ushi
of which poplulation
size is about 1 00
and the other Kuchinoshimaferal cattle in a small island, Kuchinoshima
of Kagoshima.

103

Figure 2. Genus Bubalus, Asian buffalo species.
Upper left: The swamp type of the domestic buffalo{Buablus bubalis),
male in Okinawa. Upper right: Mountain anoa {Anoa quarlesi), male. The
mountain anoa is endemic to Sulawesi of Indonesia.
The lowland anoa {Anoa depressicornis)isslightly
bigger than the mountain anoa and it also inhabits
only Sulawesi. Lower left: The river
type of the domestic buffalo (Babalus bubalis).
One of the dairy breeds of the river type, Murah
breed, is characterized
by the curly horns and back and in coat color. Lower right: The tamaraw(Bubalusmindorensis)
male. This animal is found only on Mindoro Island of Philippines.
Thebody-size and horn shape is intermediate
between the anoa and the swamp buffalo.

104

cluster.
According to the fossil record, divergence time between Bos and Synceros (orBubalus)is approximately
10 Myr (Savage and Russell, 1983). Base on this, thedivergence time between Bubalus and Synceros was calculated as 6.3 Myr (the fossil
record 5 Myr). The largest divergence within the six Bubalus species had a mean
divergence time of 2.0 Myr. All the present Bubalus species, therefore, diverged fromeach other 2.0 Myr BP or less.
Divergence time is roughly as follows; most recently 1 Myr or less between theriver type and Sri Lankan wild buffaloes,
1.5 Myr between the river and swamp typebuffaloes,
1.3 Myr between tamaraw and swamp buffalo, 2.0 Myr between anoa andother Bubalus species, and around 2.0 Myr between the two anoa species.
Molecular evolution of hemoglobin-B globins of anoa species
Kakoi et al (1994)
reported hemoglobin-P
globin amino acid sequences of
Bubalus depressicornis and B. quarlesi, and showed no substitution between the swamp
and river buffaloes, two and five between the two anoa species and the domestic buffalo,
and five between the two species of anoa, suggesting relatively
large divergence time
between the two anoa species compared to that between the two types of domestic
buffaloes. The result agrees with the cytochrome b sequence data above.
Comparison of the chromosome set of tamaraw with other bubaline species
The tamaraw had 46 chromosomes in diploidform (2N) and 58 as thefundamental number (FN) including
the sex chromosomes involved six pairs ofsubmetacentric or metacentric autosomes, 16 pairs of acroscentric autosomes and twopairs of acrocentric sex chromosomes (Namikawa et al., 1995). In contrast, all the
bubaline species of Bubalus and Synceros have FN=60 (2N=48-54)
with exception of
the swamp buffalo with FN=58 but 2N=48, previously reported elsewhere. The tamaraw
and the swamp buffalo gave unique karyotypes which may suggest a close phylogenetic
relationshipbetween them.
Source of genetic diversity of the domestic water buffalo: the river and swamptypes
The cytochrome b gene sequence analysis (Figure 6) showed that the swamp typebuffalo has a closer phylogenetic relationshipto the tamaraw {Bubalus mindorensis)than
to the river type buffalo. The wide genetic diversity in the domesticated buffalo (Figure

B

Figure3. Geographical distributionof hemoglobinin eastern Asia (Namikawa, 198 1 ).

alleles(Hbb)

of local populations(or breeds)of cattle

Figure 4. Estimated amount of gene-flow to cattle of eastern Asia, based on the hypothesisthat three-way
hybridization
had occurred in establishingprocess of the present-day cattle populations in this area.
The Pind is amount ofHbb genes originated from zebu cattle, the PNor from northern type (orhumpless cattle), and Peal from the bali cattle (see the text for the details) (Namikawa, 1981).

Figure5. Genetic distance relationshipof local breeds of eastern Asia using data from three polymorphic
protein loci and nine blood group systems of cattle. The dendrogram is by WPG clustering method(Namikawa et al , 1984).
(1) Holstein
breed of Japan, (2) Grati breed of East Java, (3) Mishima
native, Japan, (4) Tokara native, Japan, (5) Korean native, Cheju, (6) Japanese Black, (7) Japanese
Brown, (8) Korean native, Pusan, (9) Korean native, Seoul, (10) Taiwan Yellow, (1 1) Taiwan
Yellow mixed, (12) Philippine
native, Luzon, (1 3) Philippine
native, Palawan and Mindoro, (14)Philippine
native, Japan, (1 5) Padang native, Sumatra, (16) Thai native, north, (17) Thai native,
north-east, (1 9) Malaysian(West), east, (20) Aceh local breed, Sumatra, (2 1 ) Malaysian(West),
west, (22) Madura local breed, East Java, (23) Java local, Java, (24) Filial Ongole of Indonesia,(25) Local Indian Dairy of West Malaysia, (26) American Brahman of Thai land, and (27) Balicattle, Indonesia.

Figure

6. A phylogenetic
tree of the eight bovinae
method. Numerals on internal branches(Tanaka et al , unpublisheddata).

sequences constructed by using the neighbor-joining
are the bootstrap probabilities(%) from 1,000 trails

Figure7. Dendrogram drawn from the Nei's Genetic distance matrix among twenty-two water buffalo
populations,based on data of twenty five blood protein loci (Amano et al , 1992)

1087), therefore,

comes from two species or subspecies.

Conclusion
Domesticated cattle and buffaloes both have polyphyletic
origins from two or
more subspecies or species of wild bovine or bubaline.
This complexity in thedomestication
process is considered to be the primary cause of the very broad genetic
diversity in local populations (or breeds) of the domestic cattle as well as of the domestic
buffalo in the world today.
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The Phylogenetic
and DomesticatedRelationshipand Differentiation
Pigs (Sus scrofa L.) in Asiaof Wild

YAETSU KUROSAWA
Instituteof Animal Serology,

Setagaya-ku,
Present address;

Tokyo Universityof Agriculture,

Tokyo 156, Japan
Cattle Museum of Japan, Maesawa-cho, Iwate-ken 029-42, Japan

Abstract
Blood groups and biochemical
polymorphism
for the wild and domestic pigs of Asia were
investigated by the serological technique and electrophoresis
in order to clarify the phylogenetic
relationships
between them. Comparisons of erythrocyte antigens were observed. Gene frequencies for several blood
protein loci in the Asian wild and domesticated pigs were found to be very different from the European wild
pig and improved breeds. These differences were clearly observed at the Tfand Am-1 loci of serum protein.
Tf^ and Am-1Aalleles were regarded as typical alleles of the wild and domestic pigs of Asia. Theseerythrocyte
antigens and protein loci may be useful genetic markers for phylogenetic
relationshipstudies
between wild and domesticated pigs. The genetic variability
for the wild pig populations
tended to be rather
low in comparison with that of the domesticated pig population.
The analysis of genetic similarities
among
populationsof wild and domesticatedpigs substantiatedthe historicalfact that Asian and European
domesticated pigs are each derived from different wild stock.

Intro duction
Pigs (Sus scrofa L.) were domesticated from wild populationsof the same
species.
Wild pigs are distributed
throughout the Eurasian continent and surroundingislands. It has been considered that some subspecies of wild pig, for example the Indian
wild pig (Sus scrofa cristatus) and the Southeast Asian wild pig (Sus scrofa vittatus),
formed the original stock for the existing domesticated native pig of Asia. However,there have been insufficientstudies of phylogenetic
relationships
between wild anddomesticated pigs. Many questions concerning the origins of domesticated Asian pigs
remain unsolved.
The purpose of this paper is to clarify the phylogenetic
relationships
anddifferentiationof wild and domesticated pigs in Asia using information on blood groups
and biochemical polymorphism.

112

Methodology
Animals in vestigatedAccording to recent concepts in zoological taxonomy (Haltenorth,
1963), wild
pigs may be classified into about 30 subspecies, of which, the author has investigated7
subspecies and 1 6 local populations,
including wild pigs from Japan and several othercountries.
This paper focuses on blood group and biochemical polymorphism of theJapanese wild pig, in relation to Asian domesticated pig.
In Japan, there are two wild pig subspecies which may have come to Japan fromcontinental
Eurasia. The Japanese wild pig (Sus scrofa leucomystax) is found on theJapanese islands of Honshu, Shikoku and Kyushu, and the Ryukyu wild pig (Sus scrofa
riukiuanus)
is found throughout the Ryukyu islands. These islands include Amami
Oshima, Kakeroma Jima, Tokuno Shima, Okinawa Jima, Ishigaki Jima and Iriomote
Jima. There are several morphological differences between these subspecies (Figure 1).
The Ryukyu wild pig is very small in comparison with both the Japanese wild pig, andthe continental wild pig. Recently, the Ryukyu wild pig has become an endangered
species, as a result of hunting (Yasuma, 1983).
Blood typing and electrophoretic
analysis
Blood samples were collected from animals caught by local hunters in eachcountry or area investigated.
This study was completed with the help of researchers fromvarious countries including Sri Lanka, Nepal, Taiwan and Russia.
Blood typing of the wild pigs was carried out by using the blood group reagentsdeveloped for domesticated pigs. The reagents used in the study were as follows; A, O
(A system), Ea, Eb, Ed, Ee, Ef, Eg (E system), Fa, Fb (F system), Ga (G system), Ha, Hb(H system) Ka, Kb(Ksystem)
and Lh, Lk (L system). These reagents were produced by
the National Institute of Animal Industry of Japan, and they were standardizedusing theinternational
comparison test.
The genetic variation for the 22 blood proteins(Table1), each of which encodes
one locus.
These genes analyzed by means of starch and polyacrilamidegelelectrophoresis.
This analysis was performed according to the methods described by
Oishi etal (1970, 1978) and Tanakaetal(1992).

113

Table

1. List of blood protein

examined

Results

and Discussion
Blood group reagents developed for domestic pigs can be used to detect
erythrocyte antigens in wild pigs. Blood group variations in several subspecies ofSus
scrofa L., including the European wild pig (Sus scrofa scrofa), theTranscaucasianpig(Sus scrofa attila) , the Middle Asian pig (Sus scrofa nigripes) and the Far Eastern pig(Sus scrofa ussuricus) from the Eurasian continent have already been investigatedbyother workers (Buschmann, 1966; Wiatroszak, 1970; Tikhonov ^ a/., 1972), using thesamemethod of blood typing as was used for the domestic pig. The author also studied
several populations of the Taiwan wild pig (Sus scrofa taivanus) and the Indian wild pig(Sus scrofa cristatus) including the two subspecies from Japan (Kurosawa et al, 1 979,
1984,1986).

From these results, it was apparent that the characteristicof the blood groups
studied for the two Japanese wild subspecies was very different from that of the European
wild pig, and was more similar to the Asian wild subspecies.
The differences among

Figure1. Two subspecies of Japanese wild pig (adult female). Animal size is shown by the measuring rod(1 10 cm). Upper) Japanese wild pig (Sus Scrofa leucomystax). Lower) Ryukyu wild pig (Sus
scrofa riukiuanus).

Figure2. Frequency distribution
Sus scrofa L.

of the blood group antigen

Ga in different

Figure3. Frequency distribution
scrofa L.

oftransferrin

in different(Tf) alleles

subspecies

subspeciesof wild pig,of wild pig, Sus

116

Figure4. The domesticatednative pig of Nepal.

subspecies populations of the wild pig were most clearly demonstrated in the E, G, H, K
and L blood group systems. The frequency of the Ga antigen in the G system is shown(Figure2). The European wild pig characteristically
had the Ga antigen at a highfrequency, whereas some of the Asian wild pig, including the wild pig from Japan, didnot. There was a west to east geographicalcline in the frequency of this antigen. Thisphenomenon is useful for clarifying
the phylogenetic
subspeciationof Sus scrofa.
Of the 22 genetic loci of the blood proteins examined, polymorphism was foundin the Tf, Pa, Hp, Am-1, 6PGD, G6PD and ADA loci. More precisely, of the wild
subspecies studied, most of the polymorphic loci were found in wild pigs of Japan andin one subspecies fromNepal. It is particularly
noteworthy that new variants which had
not been found in the domestic pig were observed at a high frequency in each protein
locus of the Tf, Pa and 6PGD of the wild pigs from Japan (Kurosawa and Tanaka, 1988,
199 1). When comparing wild populationsof Japan with those from the continent, thegene frequencies for polymorphic loci in the Japanese wild pig were shown to be very
different from those of the European wild pig.

Figure5. Dendrogram showing genetic(UWPGM).

similarities(D)among wild and domestic

pig populations

118

The geographic distributionof allele frequencies at the Tf locus for various
populations of wild pigs in Asia and Europe is shown (Figure 3). In addition, the dataof domesticated pigs from Sumatra, Indonesia and also from Nepal which was studied
by Tanaka et al (1983a,
1992) is presented in this figure for comparison. These
domesticated pigs have similar conformation to the various wild pig types (Figure 4).
That is, although the Tf6 allele is commonly found in wild populations
from Asia andEurope, both the wild and the domesticated pigs of Asia not only have the Tf6 allele, butalso the Tf° allele. The Tf° allele, is found at a high frequency, especially,in wild
populationsof Japan at a high frequency, and moreover a new allele variant, the Tfx
allele also exists in the wild pigs of Japan. However, this was not the case with theEuropean wild populations nor the European domestic breeds (Oishi, 1 977).
Tanaka et al (1983b)
reported that the Tf° allele is widely found in Asiandomestic pig and occurs at high frequency. Therefore, it is probable that the Tf° allele
is a variant which originated in Asian wild stock. A similar result was also found for the
Am-1A allele of the Am-1 locus, that is, the Tf and Am-1 loci may be useful genetic
markers for phylogenetic
relationshipstudies between wild and domesticated pigs.
The genetic variability estimated by the proportion of polymorphic loci (Ppofy)
and the mean heterozygosity
per individual(H) was compared for populations of the
Indian wild pig, Taiwan wild pig and Japanese wild pig. The values of Ppoly and H in
the local populations of these wild pigs ranged from 0.0455 to 0.250 and from 0.0109
to 0.0884, respectively.
The genetic variability
in these wild pigs was extremely low, incomparison with that of the Asian domestic pig and other livestock.
The genetic
variabilityof Indian wild pig in Sri Lanka and the Ryukyu wild pig in Japan was thelowest, and that of the Indian wild pig in Nepal was the highest, for the wild species
investigated.
This difference in the wild pig populations
may be attributed to the effects
of isolation on island populations,whereas the continental populations
typically display
more genetic variability.
Genetic similarities
for wild and domestic pig populations
were investigatedusing genetic distance coefficients
calculated
from the allele frequencies of thepolymorphic loci according to the method described by Rogers ( 1 972). A dendrogram
which shows the genetic similarities
for wild and domestic pig populations
wasconstructed by the unweighted pair-group method (Figure 5). Several wild populations
fromAsia are similar to the domestic pig from that area, whereas the typical wild species

119of Europe are closer to the European-American breeds.
The degree of genetic
differentiation between domestic pigs and their wild ancestors for each area, i.e. Europe
or Asia, is very low. From the historicaland phylogenetic
record, Asian and European
domesticated pigs appear to have been derived from different wildstock, results in Figure
5 supports this.
Gene frequencies and thus genetic distance can be affected by mutation, natural
selection, artifical selection or geographical isolation.
As shown in the dendrogram, thedegree of genetic differentiation
among the Ryukyu wild pig populations
is quite large,
although these wild pigs are presumed to belong to the same subspecies, Sus scrofa
riukiuanus. This difference in the gene frequencies at some polymorphic loci may be dueto the effects of random genetic drift which occurred during the subspeciation
process.
In addition, populationsof this species are isolated on different islands of the Ryukyu
archipelago.
To some degree, genetic differences may reflect the complex origin or
phylogeny of each wild pig populationon the Ryukyu islands.
There are different
opinions as to whether the Ryukyu wild pig is a subspecies of Sus scrofa or a separate
species (Imaizumi 1973).
Some researchers believe wild pig constitutes
the feraldescendants of semi-domesticated pigs which originated from Southeast Asian, and was
carried from island to island by ancient peoples (Naora, 1937; Hayashida,1966). The
Ryukyu wild pigs in Ishigaki Jima or Iriomote Jima and the Southeast Asian island
domesticated pigs cluster together (Figure 5) tends to support this hypothesis.
Presently
it is not possible, to substantiate
this theory, therefore, further more detailed studies will
be required.
Acknowledgements
The author is very grateful
University
Agrobiologicalof Agriculture,

to Professor

and Dr. Takao Oishi,

Resources,

Dr. Kazue Tanaka, InstituteDepartment

and Dr. Vilen N. Tikhonov,Academy of Sciences,

Russia,for their encouragement

References
Buschmann, H. 1965

Blood group serologicalstudies.

of Animal Serology,of Genetic Resources II, National

Instituteof Cytology

and advice throughout

and Genetics,

Tokyo

Instituteofthe Russian

this work.

Proc. 9th Eur. Anim. Blood Group Conf. (Prague,1964) 131-132.
Th. 1963 Klassifikation
der Saugetiere:
Artiodactyla
1 (1 8), Handbuch der Zoologie, Walter deGruyter, Berlin.
Hayashida, S. 1960 Studies on wild boar and dog found at shell-mounds
in the Amami-Oshima archipelago.
Haltenorth,

120
J. Anthrop.

Soc. Nippon.

68:96-1

1 5 (in Japanesewith English

abstract).

Imaizumi, Y. 1973 Taxonomic study of the wild boar from the Ryukyu Island.
Jpn. Mem. Nat. Sci. Mus.
6: 1 13-129(in Japanese with English
abstract).
Kurosawa, Y., Oishi, T., Tanaka, K. and Suzuki, S. 1979 Immunogenetic
studies on wild pigs in Japan.
Anim. Blood Grps Biochem. Genet. 10:227-233.
Kurosawa, Y, Tanaka, K., Suzuki, S. and Oishi, T. 1984 Variationsof blood groups observed in wild pig
populationsin Japan. Jpn. J. Zootech. Sci. 55:209-212.
Kurosawa, Y, Tanaka, K., Nishida, T. and Cryil, H.W. 1 986 Blood groups and biochemical
polymorphisms
of wild pigs, Sus scrofa cristatus in Sri Lanka - comparison with those of other wild
populations.
Soc. Res. Native Livestock.
1 1 : 143-1 54 (in Japanese with English
abstract).

pig

Kurosawa, Y and Tanaka, K. 1 988 Electrophoretic
variants of serum transferrin
in wild pig populationsofJapan. Animal Genetics.
19:31-35.
Kurosawa, Y. and Tanaka, K. 1991 PGD variants in several wild pig populationsof East Asia. Animal
Genetics.
22:357-360.
Naora, N. 1937 On the pig in the prehistoric
age of Japan. J. Anthrop. Soc. Tokyo. 52:286-296(inOishi,
Oishi,

Oishi,

Japanese).
1977. Blood group and serum protein polymorphismin pigs and their application
as genetic
markers. Jpn. Agric. Res. Quart. 1 1:175-185.
T., Abe, T. and Mogi, K. 1970 Studies on blood groups of pigs. IV. Genetic determinationof theserum transferrin,
pre-albumin,hemopexin, ceruloplasmin
and amylase variants in pigs. Jpn. J.
Zootech. Sci. 41 :364-371(in Japanese with English
abstract).
T. and Abe, T. 1979 Polymorphism
ofPHI, 6PGD, PGD and ADA enzymes in porcine red cells.
T.

Bull. Nat. Inst. Anim. Ind. (1978)
35:9-18(in Japanese with English abstract).
Rogers, J.S. 1972 Measures of genetic similarityand genetic distance.
Univ. of Texas Publ. 72 13: 145-1 54.
Tanaka, K., Kurosawa, Y., Tomita, T. and Sihombing,
D.T.H. 1 983a Morphologicaland geneticalstudieson the native pigs in Indonesia.
Rep. Soc. Native Livestock.
10:130-139(in Japanese withEnglishabstract).
Tanaka, K., Kurosawa, Y., Doge, K., Nishida,
T. and Pradhan, S.M. 1992. Blood protein polymorphisms

Tanaka,Tikhonov,of native pigs in Nepal. Rep. Soc. Res. Native Livestock.
14:145-153(in Japanese with English
abstract).
K., Oishi, T., Kurosawa, Y. and Suzuki, S. 1983b
Genetic relationshipamong several pig
populations
in East Asia analysed by blood groups and serum protein polymorphisms.
Anim.
Blood Group Biochem. Genet. 14:191-200.
V.N., Gorelov, I.G. and Troshina, A. I. 1972 Immunogenetic
studies of wild European, Asian andAmerican Suiformes in connection with the origin of some antigens in Sus scrofa domestica.
Anim. Blood Grps Biochem. Genet. 3:173-177.

Wiatroszak,

I. 1970
Studies on blood groups in wild boar. Proc. llth Eur. Conf. Anim. Blood Grps
Biochem. Polymorphism(Warsaw, 1968).
265-270.
Yasuma, S. 1983 A critical situationof extinctionin Sus scrofa riukiuanus on Iriomote jima. Kagaku
Asahi.

43:63-67(in Japanese).

121

Questionsand answers in Session 2

Questions to Dr.Tan
Q. Did you find any amplified DNA products which are not polymorphic when you use
cattle microsatelite
primers to genotype your water buffalos?
And have you any
observations on the variation in repeat numbers? (Li)A. For details please refer to our paper in Animal Genetics 26: 355-359(1995).Briefly,of the 80 bovine microsatellites
tested on 5 buffaloes 56(70%)
gave a single PCR
product. Forty six (57%) of the microsatellites
were polymorphic. A subset of 24 locichosen based on yield and reproducibilityof the amplification
were then tested on
about 100 animals for each locus. The number of alleles per locus ranged from 2 to18.(Tan)Q. According to your D-loop data, the buffalo population from Bogor on Java clusteredwith the Murrah buffalo group. Can you explain this? (Amano)
A. Since there are 33 D-loop genotypes
found in 80 animals and we typed only 7
animals per population we attribute this anomoly to small sample size of typed
animals. (Tan)
Q. When you discuss Lankan buffaloes, do you mean wild or domesticated animals?
(M atsukawa)
A. Wecould only sample domesticatedanimals and not wild animals. (Tan)
Q. Your presentation showed clearly the genetic variation in Southeast Asian swamp
buffalo populations. Could you tell us if there is similar variation for productive traits
such as growth rate and mature body size? (Kanai)
A. Offhand I cannot quote data on such traits. However, based on morphologicalobservations there are great differences in animal sizes among animals from different
populations in the Southeast Asian region. (Tan)
Q. Lankan buffaloes are River buffaloes based upon some genetic markers, their
behaviour is, however, that of the River buffalo. Is the distinctive
behaviour of
Swamp and River buffalo a genetically
based one? (Bodo)
A. Water buffaloes wallow in order to keep cool. I presume they will wallow in whateverwater is available
when there is no choice. I have seen Lankans wallowing in clear
water. However, when given a choice Swamp buffaloes wallow in muddy water andRiver buffaloes in clear water. This is probably a learned behaviour pattern. (Tan)122

Question to Dr. Kurosawa
Q. Genetic variability
is similar between wild and domesticated forms of animals andpoultry(inthe case of ducks and chickens) and it is somewhat (20-25%) higher indomesticated forms than their ancestral wild ones in the case of quail and dogs. Whatis the situation with respect to wild pigs and domesticatedpigs?(Tanabe)
A. In Japanese wild pig (Sus scrofa lecomystax) the proportion of polymorphic loci
(Ppoly) is from 0. 1 to 0.25, heterozygosity(H) is from 0.03 to 0.08. In Ryukyu wild
pig (Sus scrofa riukiuanus) Ppoly is from 0.05 to 0. 1, H mean is from 0.01 to 0.03.
In domesticatedpig (Asian native and European-American pigs) Pploy is from 0.2
to 0.47 and H ranges from 0.08 to 0. 16.(Kurosawa)
Questions to Dr. Namikawa
Q. Hbb-X is found in Bali cattle. Is it found in any other cattle?(Astuti)
A. No breed or population has the X gene in so high a frequency as Bali cattle.
(N amikaw a)Q. How many animals per species did you sequence for cytochrome b among the
bubalines?
(Tan)
A. We only used one animal from each species or group. However, we examined as
many samples as possible using RFLP analysis, and then we selected samples forsequence study. (Namikawa)
Q. Lowland anoa and mountain anoa might be considered to be the same species. The
difference depends on their habitat. What is your opinion on the differences between
these two types? (Fukuta)
A. Molecular evolution ofb-globulinsof the lowland anoa and mountain anoa suggested
that there are large genetic differences between them. Isolation mechanisms cannot
be considered to be the cause of differentiation
between mountain and lowland anoa.
Perhaps immigration into Sulawesi took place at different times, the mountain anoa
first and then the lowland anoa. Please refer to Groves excellant work on this topic.
(N amikawa)

TECHNICALREPORTS

Session 3

Conservation

systemChairpersons
H. Inoue
S. G. Tan

Animal Genetic Resources and conservationin Vietnam

systemLE VIET LY
National

instituteof Animal Husbandry

Thuy Phuong - Tu Liem, Hanoi, Vietnam

I. Introduction:
There is considerable
diversity of animal species in Vietnam. There are 265
species of wild animals distributed
throughout the country including some rare andprecious ones. There was serious damage to wild animal life during the long war period.
According to recent evaluation by biologists
10 animal species are endangered, 18
species are vulnerable and 22 species are considered as rare. In December, 1993, thegovernment of Vietnam published"The Environment Protection Law", in which the
problem of animal genetic conservation is emphasised.
In this paper the author will discuss only domesticated animal genetic resources
cons erv ation.
II. Livestock population:
Vietnam is one of the first cradles of animal domestication.
Various species of
livestock are known to have existed in the country since the dawn of time, this include
pigs, cattle, buffalo, horses, goats, sheep, rabbits, chicken, ducks, and the muscovy duck.
The present of animal population in Vietnam is shown (Table 1).
For thousands of years, under the pressure of natural and artificial
selection
local animal populations proved to be adapted to the ecological conditions and suited tothe economic conditions of some areas of Vietnam. They possess valuable genetic
potential as they can live on low quality forage and are resistant to tropical diseases(especially parasitic diseases).
Some breeds have a reasonable reproductive performanceand produce good quality meat. Their disadvantages
is there small body size and poor
annual productivity.
III. Breeding constraints and the need to carry out the project on animal livestock
genetic conservation:
Poor breeding management as well as seasonal and irregular supplies of feed,particularlyin the dry season, are the main constraints for the rapid genetic improvement

126

of local populations.
Some indigenous breeds tend to disappear
due to low productivity
andcompetitivity
with crossbred and exotic animals.
Artificial
inseminationin pigs is extensively
practiced;
the number of
inseminated sows is around 60% of the population. Evidently, the use of semen of exotic
breeds makes the preservation of the pure native breeds difficult.
There seems to be acontradictionin the orientation towards a free marketing system and conservation ofgenetic resources. Farmers who keep local sows do not want to raise local litters because
their market price is low. Overall poor breeding management has lead to some pure
breeds becoming endangered.
The establishment and implementation of livestock conservation practices is anurgent task for the country, because it links to the development of the economy and theincome of the farmers, especiallythose who are living in marginal areas for foodproduction. They also contribute to the protection of the environment and will be usefulfor future development plans.
IV. Livestock genetic conservation programme: objectives
and measures
The main objectives of the conservation programme are:
1. To collect data of animal breeds in different ecologicalregions throughout thecountry and identify certain genes that need to be conserved.
2. To recover the economic traits of the local breeds, which are in danger, in order tokeep and improve the livestock genepool.
3. To assist the development of some objectives which have been neglected (such as
sheep for meat) or species in the process of domestication(such as Cervus Nippon
Termeric deer).
4. To contribute to economic, scientific, cultural and tourist development. The breedsof interest to the national conservation programme are:(l) Pigs:
In Vietnam the pig population is very high. In the North, Mong Cai and "I" pigs
are the two popular native stocks. The Mong Cai breed is spreading throughout country
due to a good reproductive performance, while the "I" breed is less popular as these pigs
are less prolific and tend to produce fatter carcasses. The project for the conservation ofTable 1. The present of animal populationin Vietnam

19 9 2

199 3

199 4
1 5 .5 7 0

P ie

1 2 .1 4 0

1 4 .8 0 0

C a t t le

3 .1 5 1

3 .3 3 0

3 .4 6 0

B u f f a lo

2 .8 8 5

2 .9 6 0

2 .9 7 0

G o at3 1 2 .3

3 5 3

3 70

S h e ep

Units:

D e e r (S ik a d e e r)

12

C h ic k e n

9 7 .51 0 2 .3 9 0

12
1 0 0 .6 2 7

D u ck2 7

3 1 .0 0 0

3 2 .0 0 0

fOOO heads

T pigs is a high priority

because they are still needed by the farmers of the poor humid

areas.
The mini Co pig breed still exists in some parts of the central north area of thecountry but nobody likes keeping them, due to their economic inefficiency.
The Meo and Muong Khuong pigs are mostly found in the mountainous regions.
In the south of Vietnam, the Thuoc Nhieu and Ba Xuyen are the most popular breeds.
(2) Cattle:
Vietnam Yellow cattle belongs to the humped type Asian breed with small bodysize and good hardiness.
The average body weight is 160-180kg.
There are larger
crossbred (crosses with Red Sindhi or Zebu sires ) in Vietnam. In mountainous areas
there is Muong and Moi cattle with smaller body size and strong conformation.
(3) Buffalo:
Buffalo breeds in Vietnam are of the swamp type. They are named after thegeographic name of the region where they are kept, such as Yen Bai and Tuyen Quang.
They are also sometimes named according to their size, namely, Ngo (big) buffalo or De(Small) buffalo. The average body weight is 320-350kg.
(4) Horses:
170kg.
animals.

Horses in Vietnam are all of the local type with a small body size of about 140They are mainly found in the mountainous areas and are kept as pack and riding

128(5) Goats:
Goats are of the meat type and graze mainly in the hilly rocky mountain areas.
Their development is limited.
The popular goat breed is local Co breed. The dual
purposes breed is Bach Thao has a high reproductive performance. 75% of the litters are
twin or triple. There is a clear need for a better integration of crop production, forestry
and goat grazing.
(6) Sheep:
The Phan Rang breed is the only sheep that survives in Vietnam. They belong
to the meat type. The national flock is very small (no more than 3000 heads) and islocated in Ninh Thuan province. Phan Rang sheep must be considered a priority in thelist of livestock genetic resources for conservation purposes.
(7) Chicken:
The Ri is a very popular native breed. Although it has a small body, it has a highegg yield and produces tasty meat. Several fighting chicken breeds are popular with thefarmers, as a hobby. The large meat type breeds that still exist are the Mia, Ho and Dong
Cao. Some other breeds still survive such as the Ac (black) breed, its meat is used as
tonic, the Tre breed is raised for cockfighting.
(8) Duck:
The Co duck breed is predominant and is kept in scavenger systems; they look
for their food in harvested fields. This breed is popular due to its economic efficiency(feed conversion) and good laying capacities.
The Bau duck is considered a meat type.
(9) Cervus Nippon deer:
These animals are in the process of domestication
although they still bear somewild characters and are difficult to manage and control. This deer breed is not now
endangered and is quickly spreading in the central north part of the country with the total
herd is about 12,000 heads.
(10)

Sambar deer:This animal can be seen in the zoo. But recently farmers began to create a
Sambar deer farm, in which a hundred Sambar deer are raised for meat and velvet

129

purposes.
V. Conclusion
The endangering and even extinction of domestic animals in Vietnam comesmainly from serious commercial competition with exotic breeds. With the expansion of
the artifical insemination techniques this process is occurring very quickly. The measures
to maintain genetic diversity are as follows :*Select a limited number of local breeds to be considered a first priority forconservation. These must have some economic value and be still be in demand fromfarmers in certain zones such as marginal environments. Attention must be given to the"I" pig breed, some chicken breeds such as Mia, Ho, Dong Cao and the Phan sheep.
The in situ method should be favored for animal genetic conservation. With
the renovation of the country's economy (moving towards a marketing system) we haveto contract with household-farmers
to keep the "genes" together by establishing
open
nuclear herds.
*Creating "elderly people clubs" to keep the local breeds (such as nice pet) is
an economical measure for genetic conservation.
*Preliminary studies on cryogenic preservation of semen, as well as embryotransfer in buffalo and cattle, have been conducted. These are considered an expensivemeans of ex situ conservation and are only under experimentation at this stage.
*Experiences and know-howfrom outside Vietnam is needed and international
cooperation should be considered. It involves the support in information and up to datemethodologies;
training and also equipment is needed for the ex situ methods and dataprocessing practices.
References
Do Xuan Tang.

1994.

Conservation
Le Minh Sat etal

1991.

"I" Pig

Conservationin Thanh

Hoa On the Resultsof Livestock

Genetic

in Vietnam.
On the Methodologies

of Animal genetic

Conservation.

Scientific

Informationofof Livestock

Genetic

Conservation

in

NIAH.
Le Minh Sat et al

1994.

Sika deer in Vietnam.

Vietnam. Biological
Le Viet Ly. 1990.

National

characterAnimal Livestock

Document -Ministry- On the Results

and future developmentof Agriculture

Geneticof Phan Rang sheep.

Conservation& Food Industry.

Project

in Vietnam.

Official

Project

Conservation
Philippines

Systems of Animal Genetic Resources in The

JOB M.MATIAS
College of Agriculture,

Universityof the Philippines

Los Banos College

Laguna 403 1 , Philippines
Abstract
The increasing
variationof Philippine

development
indigenous

and intensification

animal geneticof animal production

resources.

has threatened

the genetic

Concerns have been raised about the proper

systems for conservation of animals. Systems should complement the balance between human development
and animal genetic

resources conservation.

The biologicaldiversity of the Philippines
is exceptionallyrich with 2,000 fish and 960 animal
species, 500 of which are birds and 167 are mammals. Out of the 500 coral species worldwide, about 488
species in 78 genera are found in the Philippines.

Forest plant speciesof the Philippines

is one of the richestin the world with 12,000 species
Recently the objectivesof which 3,800 are endemic.
of conservation systems have been modified,from a limited

broader,

programmes.

e.g. Tamaraw (Bubalus

multi-disciplined

actionFor a particularanimalspecies,focus to amindorensis Heude), conservation efforts would include habitat conservation, protected area and community
development, conservation education and information, applied research and management, policy review andsupport activities.
This paper argues in support
genetic

resources.

conservation

An outlineof a comprehensive

is given of specificsystem approach to the conservation

animal species

conservation

to ensure rationalof animal
genetic

for present and future needs.

Introduction
In the Philippines,
there is considerable
interest in the conservation andpreservation of indigenous animal genetic resources. However, the ever-increasing meat
demand from human population has threatened the genetic identity of the native rare
breeds and strains due to the introduction of exotic temperate breeds. Introduction ofdomestic animal species fromEurope probably began in the early 1 600s, during Spanish
colonization. Some livestock and poultry from neighboring countries in Southeast Asia
and China could have entered the country centuries before.
Crossbreedingwith improved breeds is mainly based on the assumption thatindigenous breeds have low productivity.
However, clear evidence for this is lacking. The
value of native breeds in terms of sustainable production should be quantified because

132

native breeds have been exposed for many generations to the natural selection and havedeveloped adaptive mechanisms to the conditions of the tropics. This special ability tosurvive and reproduce under minimal management conditions, is a valuable attribute.
The biological diversity of the Philippines
is exceptionallyrich. At least 2,000
fish species thrive in Philippine waters. Likewise, there are 900 animal species of which
morethan 500 are birds and 167 are mammals. Approximately, 488 coral species in 78
genera are found locally out of the 500 known coral species worldwide. The Philippine
forest is one of the richest in terms of plant species totaling about 12,000 of which 3,800
are endemic. The country has a total of 61 national parks, 2 marine parks and 8 gamerefuges and bird sanctuaries. In addition, there are 1 0 wilderness areas covering an areaof 1.5 million hectares equivalent to 4% of the total land area.
It is imperative that the indigenous animal genetic resources are available topresent and future generations.
A committed effort is necessary, involving national
institutions
and individuals,
with support from the international
community to manage
these genetic resources. The systems approach to this particularundertaking should
consider the immediate need for increased animal productivity
for the expanding humanpopulation while considering the future needs for genetic diversity.
Domestic Animal Genetic Resources
Economicallyimportant poultry and livestock
species such as buffaloes(carabaos),chickens and pigs have been in the country for many centuries and were
probably domesticated fromtheir wild ancestors. In contrasts, native cattle, goats, sheep,turkey and geese are introduced animals. These animals are well adapted to local
conditions and have developed distinctive
characteristics
that merit considerations
to be
included in the country's animal genetic resources conservation program (Arboleda,
1987).

Philippine
animal production systems are still dominated by smallholders,
comprising approximately 75-80% of the total population of animals (Faylon and Roxas,
1995).
Poultry and swine predominate the industry which has shown a steadily
increasing growth rate within the last 10 years, 1984 to 1994 (Table 1). Sheep and goats
are raised as secondary source of livelihood
and animal protein. Despite this, goats haveshown a positive growth rate within the same time period, from 2.4 million heads in 1 984
to 2.6 million heads in 1994. The population of sheep is low at about 30 to 40,000

133
Table

1. Poultry,

swine, cattle,

carabao and goat inventory(in 000 heads)

Y ear

C h ick e n

D uck

S w in e

C a ttle

C a ra b a o

G o at19 8 419 8 519 8 619 8 7
19 8 819 8 919 9 019 9 11992
19 9 31994

5 9 16 1
52 3 98
5 30 06
5 324 86 032 1
6 59 126 952 86 54 80
6 3 127
6 48 68nd

5 76 152 2 152 0 852 52
5 83 36 50 172 3 682 6 88 34 08 50 68 18 77 6 127 30 47 2 75
7 03 87 5 807 90 97 9 9080 79
802 2
7 9 53
822 6

184 9
178 7
18 15
174 7
170 0
168 2
163 0
167 6
173 0
19 14
192 2

30 22
2983
2985
2 86 5
2 89 02 84 127 6 52646
2 57 6
2 57 52 55 92 36 2
2 19 1
2 17 72 0 16
2 12 022 12
2 19 2
2 12 22 30 7
2 56 2
2 63 1n d ; n o d a ta
so u rc e ; F a y lo n a n d R o x a s, 1 9 9 5 .

heads.
For large ruminants, such as cattle and carabaos, the population trend showeda decline during the past decade. In 198 1, the cattle population was around 1.9 million
heads and it decreased to 1.7 million heads in 1991, despite the increase in humanpopulation and increasing demand. However, since 1 992, it has slightly increased(Table1). The carabao population on the other hand, showed a steady decline within the last 1 0
years, from 3.0 million heads in 1984 to 2.6 million in 1994.
The Philippine
livestock development plan for a productive and sustainableindustry is anchored on specific commodity programs and an integrated approach is being
followed.
The component programs of the development plan is basicallyto upgrade thequality and quantity of the present animal resources, as follows :1.

The Breeding Cattle Development Program. The target is to increase thebeef cattle population to 3.0 million heads by 1998.
2. The Dairy Program. The introduction of 37,500 heads of dairy animals.
3. The Carabao Program. The stabilizationof the present population at 2.5
million heads.
4. The Small Ruminant Animal Program. The target is to increase thepopulation of sheep and goats to 3.4 million heads.
5. The Poultry Program. Assistance to the present industry and conservation

134
Table

2. Performance of Philippine

carabao and Murrah buffalo

P aram eters

P h ilip pin e C arabao

M u rrah B u ffalo

B irthw eig ht (kg)A d ult w eigh t (kg)M ilk yield (k g)
L actatio n leng th (d)

32
3 67
5 00
2 50

35
3 82
1 ,38 42 87

S ou rce: E u sebio , 19 80 .

T ab le 3. P erfo rm an ce reco rd of P hilippin e native p ig and B erkjala im prov ed strain
P aram eters
B irthw eig ht, k g
W eigh t at 6 m on th s,kg
W eigh t at 12 m o nths,kg
M ean age at first farrow in g,d ay s
Farrow in g in terv al,day s

Source:Penalba,

6.
7.

P h ilip pine N ative P ig

B erkjala P ig

0 .60
9 .8 534 .5
3 70 .3
2 47 .3

0 .94
16 .2069 .4
4 72 .0
2 3 1.0

1993

and improvement of indigenous chickens and native ducks.
The Swine Program. Development of new markets.
Component Programs. The stock farm development, quarantine
animal health, post production and market support programs.

stationand

The Philippine
Carabao
Recent development strategies regarding the Philippine
carabaos have involved
reproductive efficiency improvement and crossbreeding with exotic breeds from India andPakistan, and more recently, Bulgaria.
There were reports that the carabao has been
decreasing
in size and this was attributed
to the castration of bigger males geared for
draft work and that the remaining smaller carabulls were instead used for breeding(Arboleda,
1987).
The Philippine carabao (Bubalus bubalis) is classified
as a swamp-type with 48
chromosomes. The river-type buffaloes have 50 chromosomes but the crossbreeds are
fertile with 49 chromosomes (Arboleda,
1 987).
These animals are primarily used for draft purposes, both males and females, andan adult weighs about 350-400 kilograms which is relatively
smaller than that of Murrah

135

Figure1.

A female Tamaraw (Babalus mindorensis Heude) confined at the Gene pool of the Tamaraw
Conservation Project on Mindoro Island, Philippines(photo by J.M. Matias, July 1 995).

or Nili-Ravi buffaloes (Table 2). The average heightcmfor adult animals (Eusebio, 1980).
Philippine

at withers

ranges from 124 to 137

Native Pigs
Phenotypically,
native pigs in the Philippines
can be recognized by their black
hair color but occasionally, animals with white and red markings can be found. The ears
are small and erect, the snout is long, face straight,
has large eyes, trimjowls,
big neckand narrow shoulders. Characteristically,
the back is sagging with paunch belly and total
body conformation is short and narrow. Mature boars could be recognized by their tusks
but with teats. The feet are short and strong (Penalba, 1993b).
Limited available
information suggests that the pedigree of Philippine
nativepigs could have come fromnumerous species of wildpigs in the country, namely: Suscelebensis Philippinensis
Nehring on Luzon, Sus celebensis negrinus Sanborn on the
Visayas and Sus barbatus ahoenobarbus Huet on Palawan. There are indications
also
that before the Europeans arrived in the 1 600fs, pigs were introduced by Chinese traders(Penalba,
1993b).

136

Throughout the years, crossbreedingwith exotic breeds has been done and thisresulted in the emergence of strains well adapted to Philippine
conditions.
Predominantly
black-coloredstrains are called Diani and Black Ilocos, while Kaman and Koronadal
strains are colored red. Diani is the result of crossing native pigs in Batangas province
with Berkshire. Black Ilocos is the result of Berkshire, Poland China and native pig
crossbreeding.
Red Kaman is from Duroc Jersey while Koronadal was developed in
Mindanao from breeding native pigs in the region with Poland China, Berkshire andDuroc Jersey (Peiialba,
1993b).
There is the potential of native pig production in terms of additional
income forfarmers from a sustainable-typeof production system. Another point is that they areinvaluable source of genetic materials for improvement programs. The performance ofPhilippine
native pigs compared with an improved strain, Berkjala, a crossbred between
Jalajala(Luzon) native pig and Berkshire is shown (Table 3).
Philippine

Indigenous Chicken
The progenitor of the Philippine
chicken is believed to be the Red jungle fowl
{Gallus bankiva) and they were probably domesticated before written records, as earlyas the 1500's (Lambio and Gay, 1993).
The female has brownish plumage and the shank is yellow to gray and has asingle comb. The male's plumage is more attractive, with shiny red color and black tailfeathers with brown hackle. The chicken are relative small, weighing only 1.3 and 1.0
kg for males and females, respectively.
In the 1900's many exotic stock were introduced. However, there was no definite
breeding program at that time, so indiscriminate
crossing resulted in the emergence ofmanynondescript mongrels.
In 1994, out of the 64.9 million chickens in the Philippines,
72% are being raisedby smallholder farmers. The White Leghorn breed is the most popular (more than 50%of the population),and the rest are imported breeds and mongrels. The number ofPhilippine
native chicken is very small and there is no estimate of their population(Lambio and Gay, 1993).
Although the Philippine
chicken is not popular for commercial production, anumber of smallhold farmers prefer to raise them particularly
due to the very low levelof management required. In addition, they have the best mothering ability, are good

137

breeders and foragers. Health problems are almost non-existent
hardy and can survive under harsh conditions.

because they are very

Systems Approach to Conservation
The continuing depletion of biological diversity due to development has reached
an alarming state.
Efforts to conserve what is left entail a balance between theunrestricted human development and the need to preserve genetic resources.
As presented above, although indigenous breeds of animals could not compete
economically with modern-day breeds, they are important from an economic viewpoint,
and as a source of genetic Variation in breeding programs and as a national identity(Penalba, 1993a).
As a prime example, in the Philippines,
unrelenting efforts have been undertakenin the past few years to conserve the Philippine
tamaraw (Bubalus mindorensis Heude),which is a rare wildlife animal species found only on the island of Mindoro.
At present, the estimated population of the tamaraw is around 200-300 heads,
and is classified as an endangered species. In the early 1 900's, the number was estimatedto be 10,000 heads. The reasons for its decline are human encroachment, illegal logging
and the loss of grazing areas to the expanding cattle farms (Monmoiigan, 1993).
Previous conservation efforts have mainly focused on the animal itself, without
muchconcern on other factors that directly and indirectly affect its survival for the next
generations. A multi-disciplined
action programme which is at the same time integrativeand dynamic is necessary for this particular effort to succeed.
It should be recognized that the most effective way of conserving any animal
species, breed or strain and other flora and fauna is to protect its natural habitat fromfurther destruction and/or deterioration.
The deterioration
and/or destruction of thenatural habitat are, by and large, man-made(Tamaraw Conservation Program, 1 995).
The conservation program components include the following:
1.

Protected areas, community organization and community development- the focal point for this component is the people, particularly
those thatsurrounds the habitat of the conservation area. It is recognized that theirinteraction has a marked influence on the conservation efforts.

138

2. Habitat conservation- this component would include bio-physicalresources inventory (floraand fauna) and socio-economic cultural inventory, including reforestation
activities
and establishmentof community-based strategic centers.
3. Applied research- this component will handle the bio-physicaland socio-economic
characterizationof the natural habitat and to determine optimal approaches in the population restoration and/or regeneration of endangered
flora and fauna.
4. Policy studies- the review of existing laws and restrictions,ensuring in the future thepromulgations of new laws for the conservation program.
5. Conservation education, information and communications- the prime objectives of this component is to raise the level of awareness,understanding
and appreciationof genetic resources through an
expanded education and information campaign.
6. Captive species research and development- to conduct research to increase the productive and reproductive functionsof the animals.
7. Support activities- thiswould
includetrainings,
information
systems
andmonitoring/evaluationof the program.
In the interests of maintaining biologicaldiversity,
the needs of the humanpopulationand national development concerns cannot be fully sacrificed.
For theprogram to be effective, it should integrate sustainable development of animal genetic
resources which basically involved the following components :1.

2.

Accurate inventory of available animal genetic resources- this is the first step in any conservation effort wherein basic information
should be available in the form of an animal data bank.
Evaluation and documentation- the identificationof species and breeds which could be classified as
endangered or threatened is important before any conservation effort can

139

be implemented.
Appropriate conservation, in situ and/or ex situ
- the constraint of high cost is limiting the preservation of a large numberof animals but this is a very important part of the whole conservation3.

program.
4. Evaluation and use of biotechnologies- future technologies would ensure a more practicalthat needs lesser space and financial resources.

conservation

systemConclusion
Much work is still needed in order to make the present conservation efforts andsystems of animal genetic resources successful. However, the necessary technology andconcerns are gradually being developed to suit each specific species or breeds and theneeds of national development.
The urgent need is still for sufficient funds necessary for this very importantundertaking to support a comprehensive system approach in the conservation of animal
genetic resources.
References
Arboleda,
Eusebio,

C.R. 1 987.

Animal genetic

Production

Technology.

3(2):Proceedingsof SABRAO Workshop

A.N. 1980.
Oceania.

Faylon,Tropicalresources

Agriculture

Research Center,

P.S. and Roxas, D.B. 1995.
agenda of the National

Lambio, A.L. and Gay, E.C.
Technology.

conservation

The Philippine

Livestock1993.

and developmentin the Philippines.

on Animal

Resources

Animal12-15.
Genetic

in Asia andpp. 313-338.

livestockindustry:

the researchand development

Network. IRRI, Los Banos, Philippines.

Conserving

the Philippine

nativechickens.

The Tamaraw conservation

program.

Animal Production

Animal

Production

Technology.

8(1):8(1):9-13.

Momongan,V.G. 1993.

21-

23.

Penalba,

F.F.

1993a.

Agric.
Penalba,

F.F.

AnimalJ. 27(2):1993b.

genetic

resources

consortiumin RP: status

and prospects.

performanceand potential.

AnimalPhilippine

nativepigs:

Program.

1 995.

Mimeograph.

8(1):2-7.

Tamaraw Conservation

Anim. Husb. and12-15.
Production

Technology.

Collection
and Preservation
the MAFF Gene Bankof Animal Genetic Resources in

YOSHIAKI IZAIKE1 , KAZUHIRO KQCUCHI2, JUNKO NOGUCHI2, KEIJIRO MRASAWA3,
HIDEAKI TAKAHASHI3, TSUTOMU FURUKAWA3*and TAKAO OISHI2
departmentof Animal Reproduction,

National

Instituteof Animal Industry

Tsukuba Norindanchi P.O. Box 5, Tsukuba, Ibaraki 305, Japandepartment of Genetic Resources II and 3Department of Genetic Resources I,
National

Instituteof AgrobiologicalKannondai 2-1 -2, Tsukuba, Ibaraki
Present address;

Resources
305, Japan

Department of Animal Breeding

Tsukuba Norindanchi

P.O. Box 5, Tsukuba, Ibaraki

and Genetics, National

Instituteof Animal Industry

305, Japan

Abstract
organization

The MAFF genebank project on animal genetic resources in Japan was initiated in 1985. The
of MAFF genebank consists of the central bank at the National Instituteof Agrobiological

Resources and four subsidiarypreserved

as small herds,

banks.

A total of 7 1 3 breeds

cryopreserved

to preserve germplasm for the long-term to avoid inbreeding
cryopreservation
mistakes

and their lines

semen and embryos.

methods for each species,

have been collected

and are been

Cryogenic

storage has been widely adopted

depression.

The importanceand their appropriate

applicationof standardized

to prevent mishandlingandis stressed.

Introduction
Current Japanese livestock breeds and their lines can be dividing into three
categories according to their origin and history (Obata et aL , 1992). Firstly, there arevarious native livestock and domesticated fowls that either existed or were introduced
into Japan before the Meiji restoration (1 868), and some of which have kept up to thepresent time. Secondly, there are dairy cattle, pigs and poultry that were imported after
the Meiji restoration. Thirdly, there are beef cattle which are crossbreeds between nativeand imported cattle breeds after the Meiji restoration.
The increasing emphasis uponcertain breeds of livestock because of their economic value and high performance isresulting in other, less popular local breeds declining in numbers. It is recognized that
local breeds are not very useful in the modern animal industry.
However, they are
essential
for the improvement of animal breeds and research development of animal
genetic resources to maintain genetic diversity of animal resources in the country.
This paper reports on the MAFF (Ministry of Agriculture,
Forestry and142

Fisheries) gene bank project and the of cryopreservation
domesticated birds in Japan.

system of domestic animals andOrganizationof MAFF Gene Bank Project
The MAFF gene bank was initiated in 1985 for the conservation of plant,
microorganism, animal, forest species and aquatic organisms. In 1993, the DNA bankwas newly establishedand to preserve DNA and accumulate information using acomputer network. The genetic resources center of MAFF was establishedin theNational Institute of Agrobiological
Resources (NIAR) for plant, microorganism, animal
and DNA genetic resources. The MAFF gene bank consists of a national center andmanysub-banks located throughout Japan. The project aims to survey, collect, classify,
identify evaluate and preserve those genetic resources in Japan and from overseas, andfarther expand the genetic resources information stored in the database. The organizationof animal genebank is shown (Figure 1). The organization of MAFF gene bank consistsof the central bank at NIAR and 4 subsidiary banks, which are National Institute ofAnimal Industry, National Institute of Animal Health, National Institute of Sericulturaland Entomological Science, and National Livestock Breeding Center. National Livestock
Breeding Center has seven stations in different parts of Japan. The gene bank covers awide range of animals including livestock, domesticated birds, experimental animals,
silkworm and honeybees. The genebank is carried out in a collaborative
way between thecentral and subsidiarybanks.
Each member of the collaborative
team shares
responsibilities
for conservation of designated animal species.
The procedure for collection, evaluation and the category of preservation arepresented (Figure 2). When wecollect animal genetic resources, werecord passport data
which is as like a census registrationconsisting of basic information such as name of
species, breeds or line, the place and the date of the collection are entered. Evaluation of
the characteristics
of animal genetic resources is based on the animal genetic resourcescharacteristics
investigation
manual(National
Institute of Agrobiological
Resources,
1 992).
Characteristics
are classified into 3 classes (morphological,
physiologicalandeconomic), and each class consists of essential and optional descriptors.The
number ofdescriptors
varies depending on the various conditions under which animal genetic
resources are reared. An example for beef cattle is shown (Figure 2). The primarycharacteristics
of genetic resources are morphologicalcharacters such as coat color,143

presence or absence of horns. The secondary items are developmental and physiologicalcharacteristics.
The characteristics
in this category involve blood components andchromosomenumbers, and these have important implications
for the use of those animal
genetic resources practically.
The third class of characteristics
are concerned witheconomic benefits, such as age of first calving and meat production.
Preservation consist of three categories, the working, base and active collections.
The working collection means temporary preservation until the evaluation results areclear and the need for preservation is clear. However, the concept of working collection
wasdeveloped for plants, for animal genetic resources the concept of a working collection
is different.
The second category is the basic collection, which means the permanent
preservation and multiplicationof genetic resources. The third category is the active
collection consists of germplasm which can be distributed
to users.
These activities
are recorded annually in the annual reports on animal genetic
resources in the MAFF gene bank project and the annual survey report for animal genetic
resources. Textbooks and manuals for database development and cryopreservation havebeen publishedirregularly.
Statusof Collection
and Preservationin the MAFF Gene Bank
When the MAFF genebank project started in 1985 there were a total of 557
accessions preserved and the percentage of silkworm accessions represented
77%(43 1/557).
A large number of silkworm breeds and/or lines reflected Japan's industrial
background until 1950. A plan was made to collect animal genetic resources and 5 to 23
accessions have been cumulated annually. At the end of 1995, a total of 713 animal
breeds or their lines of 18 species have been preserved (Figure 3). By the year 2000 theaim is to collect and perserve about 830 accessions. The number of accessions of themain animal species, and their breakdown indicates that the number of accessions ofdomesticated birds, cattle and pigs are increasing, while silkworms have not increased.
In Japan, there are 8 native horses breeds (Hokkaido pony, Kiso pony, Taishu
pony, Nomapony, Misaki pony, Miyako pony, Tokara pony and Yonakuni pony), and2 native cattle (Mishima cattle, Kuchinosima wild cattle) and 4 crossbred beef cattle(Japanese black, Japanese brown, Japanese polled, Japanese shorthorn), and 2 native
goat (Tokara goat, Shiba goat). There are not many Japanese native animal breeds.
However, there are many domesticated native birds which consist of 17 breeds and 36

CENTRAL BANK

FigureFigure 2. The collection,

1. Organization

investigation

SUBSIDIARY

BANKS

of the MAFF animal gene bank project.

and preservation

procedure for animal genetic resoruces.

Figure 3. The annual progress

Figure 4.The number of preserved

In preservation

animal genetic

In animal genetic

resources

resources by species

in 1995.

146

Table 1.

Major animal genetic

resources preservedby the MAFF gene bank projectvarieties. The list of major animal genetic resources preserved by the MAFF gene bankis shown in Table 1. Many breeds and their lines remain to be collected.
Cryopreservationof Animal Germplasm
There are three ways to conserve animal genetic resources:( 1) in situ conservation: keeping animal population in their habitat, and reproduction isnatural;(2) ex situ preservation: animals are under artificial control for nursery and reproduction;(3) in vitro preservation: application of the technologies
of cryobiology
and reproduction
to animals.

Table 2. The probalility

ofcryopreservatinof semen and embryo in each animal species

In order to multiply endangered animal genetic resources, in situ conservation
is top priority.
In vitro preservation should be started at the same time as ex situ
preservation
is initiated.
These three ways of conservation are complementary andessential for effective conservation of animal genetic resources. In situ conservation is
the most useful method to conserve animal genetic resources. Live animal preservation
has a number of advantages. These advantages are a) cultural-historicalreasons, b) theability
to investigate
topical and new traits in the population
at any time, and c)
maintaining public awareness of the existence of the breed and interest in its frozenmaterial (Obata et al., 1994). The preservation of live animals, however, causes various
problem, for example, the high cost in keeping the animals, increased homozygosity
within small herds and depression of fertility through inbreeding.
The MAFF genebank
project aims to develop ex-situ preservation methods for animals since, in Japan, land forin-situ conservation is a constraint.
Cryopreserved germcells can be saved permanently and, not with standing any
accidents in the storage system, remain available in exactly the same condition as at thetime of their collections,at any time in future. Furthermore, cryopreservationof germ
cells and embryos could contribute not only to increase in population size but also avoid
inbreeding depression in a species. Therefore, the cryogenic storage system has been
widely developed to store sperms and embryos with a low operation cost in MAFF
genebank.
The ability to cryopreserve sperm and embryos in each of the main domestic
animals and domesticated birds is presented (Table 2). Protocols to cryopreserve sperm
has already been established.
But, the freezing of the embryo is difficult.
Only in cattle
and sheep is it practical.
In the horse, goat and pig embryos are very difficult to148

cryopreserve, so manyresearchers are engaged in developing cryopreservation
protocols
for these animals. Embryos of domesticated birds also cannot be cryopreserved, but in
1994, it was reported that cryopreservationof primordial
germ cells from chicken
embryos was successful. A chimeric bird was produced by injection of cells into anotherbreed of chicken embryos (Naito et al, 1994, Naito, 1994).
These recent successes maylead to a new means of preservation in domesticated birds.
Many methods of freezing sperm and embryo in animal species have beendeveloped.
However, the method of cryopreservationin same species needs to be
standardized
so confusion and mishandeling does not occur. There are basically4
methods to freeze of bovine embryo at the present time. These are (1) stepwise: this ismost basic method which uses 1.5 M glycerol as a freezing protectant, and after thawing,glyocerol is removed step by step out of a straw, (2) stepwise in straw: bovine embryo
is packed 1.5 M glycerol and both side are fuelled with 0.29 M sucrose at the time of
cryopreservation,
and after thawing, glycerol and sucrose are mixing by figure fillip
within a straw, (3) direct method: the bovine embryo is freezed by 1.8 M ethylene glycol,
and after thawing , it is possible
to transfer directly without any procedure, (4)vitrification:
to use a vitrification medium, bovine embryos are capable of being quickly
frozen, but after thawing, vitrification medium must be removed out of the straw step bystep. It is necessary to standardized cryopreservation
methods for each animal species
to prevent mishandling and confusion. In the MAFF genebank, a stepwise method using
glycerol has been adopted for bovine embryo freezing.
The procedure of germplasm freezing is different for each animal species.
Domesticated birds semen has specific characteristics
which make it cold temperature
tolerant within 15 minutes after ejaculation.
Therefore, ejaculated semen is collected ina test tube cooled to 5°C and after dilution with Lake solution 4 to 10 times, it is possible
to cryofreeze using evaporated liquid nitrogen immediately.
While, pig semen is very
sensitive to temperature change. The procedure for cryopreservationof pig semen iscomplex, and time consuming compared with freezing semen of domesticated birds. In
the MAFF genebank, the cryofreezing procedures for each animal species is describedin the "Animal germplasm preservation manual" (National Institue of Agrobiological
Resources, 1994). This manual describes how to get high viabilityof frozen-thawed
germplasm and lists standardized
methods and medium for each animal species. The
viabilityof frozen-thawed semen of these species varies between 50 to 90%. TheseFigure5. The procedureof pig semen cryopreservation

150

Figure 6. The percentageof accessions

preserved as live, semen and embryo in each animal species.

methods are incorporated into the stock management database.
The ratio of live and/or cryopreserved semen and embryos in the main domestic
animals and domesticated birds at the present time is shown (Figure 6). The percentageof cryopreserved germplasm in cattle and pig are 66% and 44%, respectively.
The
percentage of cryopreservationin each animal species will be increased gradually.
The
total number of cryopreserved semen tubes of cattle, pigs and domesticated birds are
23,000/0.5 ml tube, 1,400/5 ml tube and 2,500/ 0.5 ml tube, respectively(National
Institute of Agrobiological
Resources, 1995).
These cryopreserved semen have beenpreserved in the central bank and subsidiary banks.
Conclusion
Animal genetic resources that are not currently in great demand may be valuable
for scientific and economics purposes in future. When the MAFF genebank project was
initiated,
it was recommended that native animals which are endangered should becollected
and preserve first. This advisory recommendation was reasonable, but it
became clear that it is very difficult to collect a sufficient number of samples from a151highly inbreeding
endangered herd. Therefore, it is essential to collect and preserve
broadly from animal genetic resources in Japan without distinction
between native andintroduced animals. Usually, it is impossible
to build a livestock facilities
to conserve
newanimal germplasm, space for livestock is decreasing which number of accessions isincreasing.
Cryogenic systems need to be further improved. Finally, we have tocooperate harmoniously with animal genebank researchers worldwide to develop aninternational
system for animal genetic resources preservation.
References
Naito, M.,Tajima,J., Tanoue, T., Hoda, Y. and Kuwana, T. 1994.

cells in liquid
321-325.
Naito, M. 1994.

nitrogenand subsequent

production

Productionof germline

chimeric

gametes, producedby transfer

of primordial

National

Instituteof Agrobiological

Resources.

National

Institute

National

Institute

Obata,T., Takeda, H. and Oishi, T. 1994.
information.
FAO. 13:13-24.

Preservationof chickoffspring.

J. Reprod.Fertil.,of variable

chickens,

with highgerm cells.

transmission

primordial

germ
1 02 (1 1 ):rate of donor- derived

Mol. Reprod. Dev. 39: 1 53-161.

1 992. Animal genetic

resourcescharacteristics

investigation

manual for data base, pp. 192 (In Japanese).
of Agrobiological

Resources.

1 994.

Animal genetic

resources

preservation

manual, pp.

Resources.

1 995.

Annual report on animal genetic

resources.

native livestock

1 52 (In Japanese).
of Agrobiologicalgene bank project,

Obata,T.,

pp.21-38

MAFF(In Japanese).
Japanese

Takeda, H. and Izaike, Y. 1992. Preservation
Agriculture
Research Quarterly. 26 :1 -6.

of animal

breeds.

Animalgenetic

resources

genetic

resources

in Japan.

Japan

Evaluationin Japanand Preservationof Silkworm Genetic Resources

EECHI KOSEGAWA, TAKAHIRO KIKAWADA,KUNIMITSU SHIMIZU and TERUO OKAJIMALaboratoryof Genetic Resources, National

Instituteof Sericulturaland Entomological

Science

Kobuchizawa, Yamanashi 408, Japan
Abstract
In the National Instituteof Sericulturaland Entomologicalof the silkworm, Bombyx mori L., are conserved for breedingare evaluated to better

understandthe differences

Science,and experimental

among strains.

more than 450 races or strainsuse. These genetic

The traits evaluatedare classified

resources
into thefollowing three groups: (1 ) Visual characteristicsof the egg, larva, cocoon and pupa, (2) features importantin sericulture,for example hatchability
after overwintering,
rearing duration, response to artificial
diet,thermo-tolerance
biologicaland (3) the cocoon features in relation

characterssilkworm geneticof the silkworm.

to reeling

Recent topics in relation

and both biochemical
to the evaluationand molecularand preservationofresources in Japan will be discussed.

Domestic silkworms in Japan
In Japan, races of domestic silkworm, Bombyx mori L., can be categorized into
several groups based on their uses. All races for Japanese cocoon production are"Authorized
Races" they are Fi hybrids whose parent races and mode of crosscombination were authorized by the government. Cocoon production using unauthorizedraces is prohibitedby the Sericultural
Industry Law of Japan. This law was enacted tostandardize raw silk quality using Authorized Races, and the breeding of silkworm races
suitablefor mechanized reeling of silk.
Authorized Races were bred from other
Authorized Races or unauthorized silkworm races conserved solely for breeding. Various
breeding stocks have been conserved by many breeders. Mutant strains have beenconserved for basic science as well as applied biology.
Silkworm mutants havecontributed greatly to classical genetics in Japan (Tanaka, 1927).
Domesticated silkworm cannot survive in natural environments since the larval
body is too heavy to move on a mulberry branch to find the leaves. Also domesticatedlarvae have no mimicry to escape predators, in particular
birds, and the adults havemating difficulties.
However the wild ancestor of the domesticated silkworm has notclearly been determined. A possible ancestor is Bombyx Mandarina Moore, which has
not been used for cocoon production (Yoshitake,1984).
B. Mandarina can produce

154

fertile hybrid progeny when mated with domestic silkworm. Strains
hybridization
using B. Mandarina are conserved in several institutes.

derivedfrom

Institutions
conserving silkworm stocks
National Institute of Sericulturaland Entomological Science
National Institute of Sericultural and Entomological
Science (NISES) conserves
many silkworm races for a variety of uses. Many stocks have been used to produce
Authorized Races. Stocks used to produce Authorized Races are conserved for future
breeding purposes and continue to be used for scientific studies even after governmental
authorization
has been terminated.
Some silkworm races have been maintained morethan 200 years in Japan. The silkworm races and strains conserved at NISES are divided
into 5 groups: a) Authorized Races b) Geographically
fixed races c) Improved races d)
Breeding resources e) Mutants. Groups b, c and e are categorized as "genetic resourcesof NISES", whereas groups a and d are not included in this category (Sorita, 199 1). The
genetic resources of NISES are distributedfor both research and breeding.
Kyushu University Institute of Genetic Resources
Kyushu University Institute of Genetic Resources conserves more than 400
mutant strains of domestic silkworm. The mutant stocks have been used in basic
genetics, physiology,
biochemistry
and molecular biology.
All stocks in Kyushu
University are multi-marker strains each containing several distinctive
mutant genes
located on different chromosomes. The multi-marker strains are being used to establish
a linkage map of B. mori (Doira, 1978) and are offered to investigators
on request. The
silkworm is an important experimentalorganism like the fruit fly, Drosophilamelanogaster.
Private companies and an incorporated organization
Some private companies and an incorporated organization
have originally bred Authorized Races and stocks for breeding.

relatedto sericulture

Silkworm genetic resources in NISES
The genetic resources of the silkworm at the NISES are conserved in the
Laboratory of Genetic Resources, Kobuchizawa, Yamanashi Prefecture. These genetic
resources are classified as listed in Table 1.

Table1.

The number ofgeographicalraces of domestic
Sericulturaland Entomological
Science

Table 2a. The items to be checked

during the conservation

silkworm stocked in Nationalof silkworm stocks (Primary

Instituteofcharacters)

Geographic races
The category of geographic races conserved in NISES involve four groups,
Japanese, Chinese, European and Tropical races. Three groups, Japanese, Chinese, andEuropean races, have contributed to cocoon production in Japan. Tropical races wereused to establish lines that can tolerate both high temperature and humidity. Each groupis classified
into, native races and improved varieties.
Native races had been used forJapanese sericulture before the inception of the authorization
law. These races are
unsuitable for the production of raw silk of standardized
quality, since they have defects
in the mass production and in the raw silk percentage of the cocoon. Native races areused to improve and stimulate breeding resources and materials for experimental

Table 2b.

The items to be checked

during conservation

Table 2c.

The items to be checked during conservationof silkworm

stocks (Secondaryof silkworm stocks (Tertiary

characters)

characters)

purposes. Improved races, once authorized by the government, are used in sericulture as
Authorized Races. Inter-crossing of geographic groups is an efficient way to obtainheterosis.
Moltinism

races
Most silkworms are tetramolters, however, races from Korea vary in the numberof larval moltings and are classified
as moltinism varieties.
There are trimolters,
pentamolters and one hexamolter in races from Korea. Some trimolters are used insericulture since these produce cocoons with a particularly
fine filament. The raw silk
produced by such races is suitable for high class silk products.
Mutant strains
Mutant strains are divided into original mutant strains with one mutant gene andmulti-marker strains (most of the later have come from Kyushu University).
Someoriginal strains have contributed to experiments on breeding of commercial races, for157

example oligophagous races that can be reared on an artificial
diet, and races possessing
sex linked body markings. Mutant genes preserved in these strains serve as usefulgenetic markers in, for example, linkage analysis and three point analyses (Doira, 1978).

Conservation of silkworm stocks in NISESThe conservation of geographic races and moltinism races, the maintenance ofgenetic diversity is important because of the possibility
that these races possess unknown
heterogeneous characters, in addition, to the known practical characters. Usually, fifty
or more moths are used for egg production to avoid the loss of minor genes from the
population.
In this way races are conserved as population gene pools.
Authorized races and breeding resources are maintained under high selection
pressure on the basis of phenotypes related to raw silk production. Consequently, theseraces can produce silk of good quality. However, this type of selection is not always
performed whenconserving silkworm genetic resources. Consequently, silkworm genetic
resources are not in the best state for cocoon production.
Mutant strains are maintained
only for the preservation of genes or genotypes. Although the technology of the rearing
silkworm on an artificial
diet has been establishedand generally applied to thecommercial production of cocoons in Japan, an artificial
diet is not used to maintaingenetic resources. This is because the silkworm is considered to be adapted only tomulberry leaves and there is the fear that dietary selection would occur if silkworms arereared on an artificial diet.
Preservation of diapause eggs
Each race is reared once a year except for the tropical races which are reared
twice or moreper year. In winter, silkworms are preserved as hibernating diapause eggs,
since mulberry leaves can only be produced in southern Kyushu and Okinawa during
winter. Rearing seasons are limited to when mulberry leaves are available.
We havethree major rearing seasons in Japan, spring, summerand autumn. The winter diapause
stage enables silkworm eggs to survive low temperatures and low relative humidity. For
the efficient preservation of silkworm eggs, the diapause phenomenon of silkworm eggs
should be understood well.
The diapause in silkworm eggs is induced by the action of the diapause hormone,

158

which is secreted from the maternal suboesophageal
ganglion and influences the ovary.
The newly laid eggs enter diapause at the early gastrula stage. The eggs develop again
after longterm chilling.
The induction of diapause is also geneticallycontrolled byvoltinism.
In univoltine races, moths always produce diapause eggs. The diapause ofboth bivoltine races and polyvoltine races is determined by temperature and photoperiod.
In Japanese sericulture, the diapause phenomenon in most races and strains of thedomestic silkworm is well controlled by artificial procedures. To induce diapause eggs,
the eggs of the parental generation were incubated at 25°C under long day photoperiod,
15 hours photophase and 9 hours scotophase. As a result, the female moths produce eggs
that enter diapause.
The diapause eggs are preserved at natural temperatures until
autumn and then transferred to a low temperature until the following spring. Diapause
is terminatedunder cold temperatures during winter. Diapause can be broken by
subjecting eggs to hot hydrochloric
acid treatment. The hot acid treatment is effective
against the eggs for a few days after oviposition
or after chilling for more than one
month. If eggs have already been kept cold for more than 1 50 days, then the eggs shouldbe transferred to 25°C without acid treatment. Thus the life cycle of silkworm can becompletely controlled.
Diapause eggs are difficult to obtain from tropical races in Japanese conditions.
Therefore the larvae of these races are reared at relatively low temperatures and alteredphotoperiod. Under these rearing conditions, an adequate number of diapause eggs are
obtained. However, the eggs are intolerant of preservation by longterm chilling.
Thustropical races are reared twice, in spring and autumn, in Japan.
In the case of diapause mutants, the homozygotes develop without diapause eventhough the eggs have received diapause hormone. The mutant strains are conserved asheterozygotes that can enter diapause.
Systematic

rearing of silkworms
During a rearing cycle, for efficiency,
silkworm stocks should be reared
synchronously.
Synchronous rearing minimizes accidental loss of genetic resources.
Much information useful for the planning of synchronous rearing under standard
conditions
has been accumulated, for example, the larval life span, hatchability
andmating combination. In tetramolters and one pentamolter that have longer life span at thelarval stage, the starting day of rearing is adjusted to a few days earlier than in other159

races.
Error management during conservation
Although there is no way to completely prevent errors during conservation, we
can predict which errors are likely to occur most frequently. Therefore several strategies
have been designed to minimize errors.
For all races, 50 batches of eggs, each batch from one of 50 moths, are firstprocessed for rearing. The additional eggs are stored at a low temperature, and can beused for additional rearing if the first rearing fails. Chemical disinfection
and washingof rearing rooms and instruments after the end of each rearing period is important toprevent diseases. Pebrin caused by a protozoa is the most serious disease because of thetransovarian transmission. Therefore, we are obliged to inspect female moths for pebrin
according to Sericultural
Industry Law. In addition, good maintenance of refrigerator
machinery is essential for egg preservation.
To prevent the loss of silkworm stocks, theeggs of each batch, each produced by one moth, are divided into two groups which are
kept in different laboratories.
The location of rearing is an important factor in theconservation of silkworm stocks. In the summer season in Kobuchizawa Town, theaverage air temperature of around 25°C, relative humidity is low and good quality
mulberry leaves available which make silkworms healthy.
Characteristics
of silkworm genetic resources
Each race of silkworms is phenotypicallydistinguishable.
carefully examined during each rearing period. The subjects for checking
as follows.

All stocks areare summarized

Visually

distinctive
characters
Visually distinctive
characters are used to identify races and sometimes toindicate the occurrence of contamination.
In some cases, genes which control visualcharacters are known and the location on specific chromosomes has been determined.
Primary visual characters which are always recorded are listed (Table 2a).
Physiologicalcharacters
Even whenthe responsible genes are not understood, physiologicalcharactersinherited as prominent features, typical of each race, for example, the tolerance against

cDNA

160

extreme temperatures, are recorded. Such features and related ones needed for theconservation of silkworm stocks are considered secondary traits (Table 2b).
Features important for raw silk production
Most of the races of silkworm were or are used for cocoon production.
Therefore, the features which are related to reeling of silk are important and recorded
periodically.
These characteristics
are considered tertiary evaluation characters (Table
2c).

Biochemical evaluation
The number of biochemical traits recorded for silkworm stocks is limited.
However, several proteins and enzymes have been analyzed for their location on
chromosomes and the frequency of allelic genes among the silkworm stocks. These data
are sometimes important to identify the stocks for molecular markers, for example
amylase (Matsumura, 195 1) phosphatase(Yoshitake
and Akiyama, 1965), esterase(Eguchi et al, 1965; Eguchi and Yoshitake, 1966; Yoshitake et al, 1966), proteinase(Eguchi and Yoshitake, 1967), proteinase inhibitors(Fujii et al, 1989; Eguchi andKanbe, 1982), haemolymph proteins (Gamo and Ohtsuka, 1 980; Kawaguchi et al, 1970;
Obara and Watanabe, 1969), storage proteins (Tojo et al, 1980), yolk proteins (Izumi
etal, 1980; Me and Yamashita, 1983).
Pathologicalevaluation
The diseases of silkworm have been investigatedin NISES and in otherlaboratories.
The susceptibilityof silkworm to several viruses has been investigatedinmost of the silkworm genetic resources (Furuta, 1995). A noteworthy result is the recent
establishment of a silkworm race which is resistant to densonucleosis
virus (Eguchi et al,1991).

Molecular evaluation
Molecular techniques applied to genomic andclones of the silkworm
should be undertaken to obtain new perspectives on genetic resources.
One maintechnique for the silkworm is Southern blot analysis. Genomic clones constructed for thistype of analysis (Yukuhiro et al, 1993; Tamura et al, 1993) were found to haverepetitive
sequences or unique sequences. Two races, C108 and p50, which are161

commonly used as the standard silkworm races for genome study, were revealed to be
cDNAdistinctive
by Southern analysis using such genomic clones. Alsoprobes wereconstructed from poly(A)+ RNAs of silkworm eggs and used for Southern analysis(Hara, 1996).
Some of the probes could detect inter-race differences,although otherprobes exhibited neither inter- nor intra-race differences.
The polymerase chain reaction(PCR) methodology has been applied to most of the geographic races stocked in NISES(Yasukochi,
personal communication)
this has uncovered Randomly Amplified
Polymorphic
DNA (RAPD).
These techniques
are helping clarify phylogenetic
relationships
among the races.
Conclusion
Currently the silkworm is being used in new areas, for example, for medical uses
such as producing fibroin membranes. Consequently, the evaluation of silkworm genetic
resources should be updated to respond to newneeds. The genetic stocks of the silkworm
in Japan exceeds 1,000 accessions.
The establishmentof systematic and authorized
information system on these stocks is necessary.
References
Doira, H. 1978.

Genetic

stocks of the silkworm.

- The silkworm:

an importantlaboratory

tool.

Kddansha

Japan,53-81.
Eguchi,M. and M. Kanbe 1982 Changes

in haemolymph

silkworm, Bombyx mori L. (Lepidoptera:protease

Bombycidae).

inhibitors

duringAppliedEntomology

metamorphosis
and Zoologyof theJpn.

17:179-187.
Eguchi,M., N. Yoshitakeand H. Kai (1965)

Eguchi,M. and N. Yoshitake

Eguchi,M. and N. Yoshitake

Eguchi,R, O. Ninaki and W. Hara 1991 Genetical

Bombyx mori L. Jpn. J. Genet.
1966 Genetic

Bombyx mori L. Jpn. J. Genet.
silkworm,

1967

H. H. Aratake,characterization

Furuta,Y. 1995.

of blood

esterasein the silkworm,

on isozymes of the integument

esterasein the silkworm,

41: 267-273.

Electrophoretic

J. Seric.

variations

ofproteinaseof a novel chymotrypsin
hemolymph

in the digestivejuiceof theto densonucleosis

virus

1989

and

843-844.
analysisSci.

on the nonsusceptibility
Jpn.

60: 384-389.

D. L. Rong, M. Nakamura, Y. Kawaguchi,

act-A) gene from the larval
å Physiol.

and inheritance

40: 15-19.
studies

Bombyx mori L. Nature 214:in the silkworm,Bombyxmori.
Fujii,

Types

inhibitor

B. Sakaguchi

controlledof the silkworm,

Purificationby the chymotrypsin

inhibitor

A

Bombyx mori. Comp. Biochem.

andB. 94: 149-155.
Susceptibilityof the races of the silkworm,

Bombyx mori, preserved

in NISES

to the162
nuclear polyhedrosis

virus and densonucleosis

virus.

Bull.

Natl.

Inst.

Seric.

Entomol.

Sci.

15:

Gamo, T., and Y. Ohtsuka (1980)
Phylogenetic
studieson the racial differentiationof the basis
polymorphic
genes in haemolymph proteins.
Bull. Sericul.
Exp. Sta. 28: 15-50.

of

119-145.

Hara, W. 1996 Linkage
Natl.
lrie,

analysis

Inst.

Seric.

K., and O. Yamashita
proterties,

and identificationof strain

by RFLP in the silkworm

Entomol.

Sci.

(1983)

Egg-specificproteinin the silkworm,

and titre changes

duringoogenesis

localization

Bombyx mori. Bull.

16: In press,
Bombyx mori: Purification,and embryogenesis.

InsectBiochem.

13:71-80.
Kawaguchi,

Y., H. Doira and H. Chikushi
Proc. Sericult.

Sci.

Kyushu.

Matsumura, H., 195 1 Distribution(1970)

Kaikotaieki-tanpakushitsu

no idengaku-teki-kenkyu.

1: 5 1. (in Japanese)of the amylase

type in various races of the sikworm, Bombyx mori L. 1 3:

521-533.
Obara, R., and H. Watanabe (1 969)
BombyxmoriL.
Sorita, S. (1991 ) Kaiko-iden-shigen

Science,

Agarose-gel

J. Seric.

Japan.

Sci.

(1993)

GammaField

Y. Yasukochi,

RFLP patterns
Symposia,

detected

Yoshitake,of Sericultural

N. Hinomoto, K. Shimizu,

by the transposons,Japan.

1-4. 1 Thirdin the silkworm,
and Entomological

E. Kosegawa and T.

MAG and Kl.4

in silkworm

races.

(1980)

Storageproteins

edition(in Japnaese).

in the silkworm,

Bombyx mori. Insect

10:289-303.

N., and M. Akiyama 1 965 Distribution
K., Yasukouchi,
silkworm,Bombyx(1C13).1.

Institute

Meibundoof the silkworm,BombyxmoriL.
Yukuhiro,

proteins

NIAR MAFF Japan. 32: 47-62.

S., M. Nagata and M. Kobayashi
Biochem.

of hemolymph

no hozon to riyo. National

Tanaka, Y. 1927 Kaiko no iden-kowa.
Tojo,electrophoresis
38: 386-394.

1-68 (in Japanese).

Tamura, T., T. Kanda, K. Yukuhiro,
OkajimaJpn.

of types

J. Seric.

Sci.

of the blood add-phosphatase
Jpn.

Y., Kanda, T. and Tamura, T.
mori, efficiency

of collecting

in various

strains

34: 99-103.
1993

Construction

probe to detect

RFLP.

of RFLP map of the
Jpn.

J. Genet.

68: 448

163

Questionsand answers in Session 3

Questions to Dr. Matias
Q. Why do people like black pigs in the Philippines?
(Wang)
A. In fact white pigs are more popular in the Philippines
e.g. Landrace and Yorkshire
breeds. We have in the Philippines
a special dish for pigs, that is, roasting the whole
pig, wherein white colored pigs are prefered. However in some speciality shops native
black pigs command a higher price than white pigs. (Matias)
Q. What is the productivity of pigs in the Philippines,
for example what is the slaughter
weight? (Wang)
A. The productivityof native pigs is not as high as exotic pigs or crossbreds. The
slaughter weight for native pigs (in my experience from the Ifugao) ranges from 3040kg. (Matias)
Q. According to our direct observations the surviving number of Tamaraw is only 43.
However the count is continuing.
Do you have any comment, Dr. matias?
(N amikawa)
A. Unofficial reports suggest the population of the Tamaraw is 200, but the exact data
is not available. (Matias)
Q. Do you have any idea what the breeding structure of the Tamaraw is? (Namikawa)A. Tamaraws are very secretive and shy animals. Efforts are still being made to
determine such factors. (Matias)
Q. Would one reason for the decrease in size of the carabao be that farmers sell off thelarger animals to get a better price? Leaving only small animals for breeding? (Tan)
A. I agree with your observation. Another reason for the decline in size is that farmers
castrate larger bulls for draft purposes. Smaller animals are used for natural matingwith females. (Matias)
Q. Do you have any evidence or historical stories concerning the domesticationof native
chickens or pigs in the Philippines?
(Yamamoto)
A. We do have written reports regarding domesticationof native chickens and pigs.
Howeverscientic facts should be established first before we can resolve issues related
to domestication.
(Matias)Question to Dr. Izaike
Q. In your genebank there are 33 breeds/linesof pig. How many are breeds and how
many are lines or strains? Where are they from? (Wang)
A. We collected 5 breeds the rest are lines. The breeds came from Europe and China.
(Izaike)TECHNICALREPORTS

SessionNewtechnology

4for conservation

Chairpersons
Y. Kanai
K. Sekikawa

Use of Marker Informationin Small Populations

to Maintain

Variability

HISATO TAKEDA1 *, TARO OBATA1 , TSUTOMU FURUKAWA2#?
KEIJIRONIRASAWA2 , HIDEAKI TAKAHASHI2

departmentof Animal Breeding and Genetics, National Instituteof Animal Industry
Tsukuba Norindanchi
P.O. Box 5, Tsukuba, Ibaraki 305, Japan
2Department of Genetic Resources I, National Instituteof Agrobiological
Resources
Kannondai 2-1 -2, Tsukuba, Ibaraki 305, Japan*Present address: Department of Genetic Resources I, National Instituteof Agrobiological
Resources
Kannondai 2-1 -2, Tsukuba, Ibaraki 305, Japan^Present address:
Department of Animal Breeding and Genetics, National Institute of Animal Industry
Tsukuba Norindanchi
P.O. Box 5, Tsukuba, Ibaraki 305, Japan

Introduction
It is essential
to keep genetic variability
as high as possible in order to avoid
inbreeding
depression
or disappearanceof valuable genes, when we keep animal
resources in a small population. We usually use coefficientsof inbreeding, relationship(Wright, 192 1) or coancestry (Malecot, 1 948) as criteria to measure genetic variability.
Coancestry is the correspondence probability
that a pair of gametes in the
population inherited from the same gamete of the same ancestry, and other coefficientscan be derived from coancestry. The calculation of this probabilities
based on the ideathat offspring inherit one gamete from each parent. Without information about whichof the pair is inherited, we can only give probabilityof 1/2 for each (Falconer, 1960).
Figure 1 shows a simulated distributionof proportion that the gamete of X andY inherited the same grandparental
allele at 1000 loci in the family of Figure 2.
Coancestry gives us only one value of probability,
0. 125, for this proportion in thisfamily, while the simulated values are widely distributed.
That means, more precise
evaluation is possible with some additional information.
Fortunately, many loci on animal genome maps have been located and where wecan distinguish
the genotype of each individual
exactly (Cheng et ah, 1 995 ; Archibald
etal, 1995; Crawford et al\ 1994; Bishop et al, 1994; Rohrer et aL, 1994; Bumsteadand Palyga, 1992).
The genotype of parents and offspring sometimes gives us
information about which of the parents or which gamete of the pair in the parent, thegamete of the offspring is inherited from.
Wecan make moreprecise evaluation of coancestry using information on theseloci as markers.

168

MethodWithout information about marker genotypes, the probability
that one gameteof an individual X and another individual
Y have the same origin, r(X,Y) is the averageof the probability
that the gametes of Y and the parents having the same origin,r(X,Y)=

l/2 {r(P,Y)+r(M,Y)}.

While, if sire(P),dam(M) and offspring(X)have genotype of AB, AC and AC
at one marker locus, we can easily know that the allele A of the offspring is inherited
fromthe sire and C from the dam. Further, at the locus which have recombination ratio
c with this marker locus, the allele on the gamete having A inherited from the paternal
gamete having A with probability(1-c), and from the paternal gamete having B withprobabilityc. Then we can evaluate the probabilityof correspondence at this locus as,r(XA,Y)=(l -c) - r(PA?Y)+c- r(PB?Y).

where, subscript A and B denotes the allele on the gamete having marker allele A and B,
respectively.
Table 1 shows the calculation of the correspondence probability
for eachcombination of marker genotypes of parents and offspring.
Allele type A, B, C, D
denotes only the pattern in this table, and it covers all the possible cases. Then some
generations
after the base population,
we have the probabilityof correspondence
expressed as a polynomial of the recombination rate c.
Now,wehave a method to evaluate the probability
that a pair of the gametes arecorrespondent at one position around a marker locus as a function of the recombination
ratio, f(c).
Then integral of the function and accumulation for all the marker lociscattered on the gamete leads us to the average probability
for whole gamete,r=£J wf(c)dc
where wdenotes the weight for each position

and£Jwdc=l.
Weuse ordinary coancestiy instead of f(c) for the area without available marker. Usually,
wis the same for all positions
on the gamete. We can, however, put other values for
position with special importance, such as QTL or MHC (Takeda et al.9 1995).

Table1. Calculationof the correspondence

probabilityof offspring

The probability
that one gamete of offspring and other gamete(Y) in the population
inherit allele ofsame origin at locus of recombination ratio c with the marker, r(Xi,Y), can be calculated from theprobability
between parental gamete and Y, r(Vj,Y) and r(Wk,Y). Subscript denote the type of marker
allele that the gamete have. This table covers all the possible cases, since allele type A, B, C, D and theparents V ,W denote only the pattern.

Simulation
A simulation study was conducted for this method of evaluating correspondence
probability(Takeda et al 1996). It was assumed that a gamete have 10 linkage groups
and each has a length of 100 cM. It was simulated whether the gamete chosen from X

170
Table 2. Mean, standard deviation of evaluated probability
and correlation between evaluatedand realized proportion with different number of available marker locicasenumber of, , .
marker loci

mean

,
s.d.

,..
correlation

1
2
3
4

10
20
30
40

0.12510.12500.12510.1251

0.017
0.026
0.030
0.030

0.474
0.599
0.668
0.745

The distributionof evaluatedprobability
larger. The correlation
becomes higher
Table

probability

becomes wider as the area covered with markers becomesas the number of available
marker becomes larger.

3. Mean, standard deviationof evaluatedand realized proportion
with different

number of, .
marker locihetero.^zygosity

2
3
4
5

0.50
0.67
0.75
O80

,

The probabilities
number of allele
heterozygosity.

åå

probability
and correlation
between evaluatednumber of allele types on each marker locus

probability

mean

,
s.d.

, +.
correlation

0.12500.12510.12510.1250

0.018
0.029
0.036
0.040

0.584
0.745
0.815
0.850

were calculated
for different
numbers of allele types of each marker in case 4. As thetype loci becomes larger, the marker becomes more informative
because of the higher

and Y in Figure 2 inherited the same grandparental allele on a loci at every 1 cM (total
1000 loci). The proportion of loci where inherit the same allele, realized proportion, wascompared with the evaluated probabilityby this method.
The evaluated probabilities
were calculated for 4 cases having different numbersof available marker loci (Figure 3). Case 1: 1 marker located at 50 cM from one end ofeach linkage group (total 10 markers) was used to evaluate the area from 30 to 70 cM.
They covered 40% of whole gamete and coancestry 0. 125 was used to other area. Case2: 2 markers located at 30 and 70 cM of each linkage group (total 20 markers) were used
to evaluate the area from 10 to 90 cM. They covered 80% of the gamete. Case 3: 3
markers located at 20, 50 and 80 cM of each linkage group (total 30 markers) covered
the whole gamete. Case 4: 5 markers located at 10, 30, 50, 70 and 90 cM of each linkage
group (total 50 markers) covered the whole gamete.
Three types were randomly put on the grandparental
allele of each marker locus.
Then, we change the number ofallele types put on each marker locus from 2 to 5 in Case
4. These evaluations were done for 100,000 simulated families.

Figure1. The simulated distributionof the proportion that the gametes of X and Y of family in Fig. 2
inherited the same grand parental allele. The proportion has mean 0. 125 as coancestry indicate,but it is distributed
from 0 to 0.50 and has a deviation of0.067.

Figure 2. The family of the simulation.
X and Y are cousins each other both paternal and maternal. It issimulated
whether a pair of gametes from X and Y, one from each, inherit the same
grandparental
allele on loci throughout the gamete.

Figure 3.

Position of the markers and the area where the probability
is calculatedusing markers for eachcase of the simulation.
Each of 10 linkage groups has markers at the same position. Case 1 :1 marker located at 50 cM from one end of each linkage group was used to evaluate the areafrom 30 to 70 cM. The coancestry 0.125 was used for the other area. Case 2: 2 markers locatedat 30 and 70 cm of each linkage group were used to evaluate the area from 10 to 50 cm andfrom 50 to 90 cM respectively. Case 3: 3 markers located at 20, 50 and 80 cM of each linkage
group were used to evaluate the area from 0 to 35, 35 to 65 and 65 to 100 cm respectively.
Case 4: 5 markers located at 10, 30, 50, 70 and 90 cm of each linkage group were used toevaluate the area from 0 to 20, 20 to 40, 40 to 60, 60 to 80 and 80 to 100 cm respectively.

Result and Discussion
A distribution of realized values simulated has already shown in Figure 1. The
meanwas 0.125 as coancestry indicate, but it ranges from 0 to 0.50 with standard
deviation of 0.067. Figure 4 shows a simulated distributionof the evaluated probabilityof case 4. Figure 5 shows the scattered diagram of the evaluated value of case 4 and therealized value of the first 1000 simulations.
Table 2 shows the mean and standard
deviation of evaluated probability
and the correlation with realized proportion.
The distribution of evaluated probability becomes wider as the area covered withmarkers becomes larger. The correlation becomes higher up to 0.75 in case 4 as thenumber of available marker becomes larger. These correlations are high even 0.47 of173

Figure4.

Figure 5.

The simulated
The distribution

Scatteredproportion

distributionof the evaluated probability.
The probability
calculated
for case 4.
ranges 0.03 to 0.30 and has mean 0.125, standard deviation 0.03.

diagram of the evaluated probability
and the realized proportion.
The realized
and the evaluated probabilityof the first 1000 simulations in case 4 are plotted.

174case 1, since the ordinary coancestry have only one value of evaluated probability,
0. 125.
Table 3 shows the mean and standard deviation of evaluated probability
and thecorrelation
between evaluated probability
and realized proportion in case 4 when wechange the number of allele types on each marker locus. The marker becomes more
informative because of the higher heterozygosity
as the number of allele types becomeslarger. The correlation reached 0.85 with 5 types of alleles on each marker loci.
Although the situations in this simulation, the number and length of linkage
group or the position of marker loci, are different from that in real animals, the value ofcorrelation would not be so different, if the heterozygosityof markers and the ratio of
area covered are similar.
This simulation indicated that use of marker information gives us more precise
evaluation for the relationship between gametes than the ordinary methods. It enables us
to select the mating pair effectively to keep genetic variability
high in a small population.
Reference
Archibald,

A.L., Halay, C.S., Brown J.F. et al
scrofa).

Bishop,

1995.

The PIGMaP consortium

linkagemap of pig (Sus

Mammalian Genome 6: 1 57-175.

M.D., Kappes,

S.M., Keele,

J.W. et. al.

1994.

A Genetic

Linkage

Map for Cattle.

Genetics

136:619-639.
Bumstead,N. and Palyga,

J.

1992.

A Preliminary

Linkage

Map of the Chicken

Genome. Genomics

13:690-697.
Cheng,

H.H., Levin, I., Vallejo,

R.L. et al.

Markers of High Utility.

Poultry

1995.

Development

Science74: 1 855-1

Crawford, A.M.,Montgomery, G.W.,Pierson, C.A. et al.
Groups Derived

From the Analysis

Falconer,

D.S. 1 960.

Introduction

Malecot,

G. 1948.

Les Mathematiques

1994.

of Paternal

to Quantitativede l'Heredite,

Map of the Chicken

with

874.
Sheep Linkage

Half-Sib

Genetics,of Genetic

Families.

Mapping:

Nineteen

Genetics

137:573-579.

Oliverand Boyd, Edinburgh.

Mathematics

of Heredity(EnglishLinkagetranslation),

Freeman, SanFrancisco.
Rohrer,G.A., Alexander,

L.J., Keele, J.W., Smith T.P. and Beattie

Map of the Porcine

Genome. Genetics

Takeda, H., Obata, T. and Satoh, M. 1995.
(in Japanese),

Proceedings

Genetic

Relationship

Use of Geneticof 90th annual

Takeda, H., Obata, T., Satoh, M. and Togashi.
(in Japanese),meetings

K. 1996.
Proceedings

Sci.:181.
Wright,S. 1921.

Systemof mating.

Genetics

C.W. 1994.

A Microsatellite

Linkage

136:231-245.

6:1 ll-178.

Markers to Evaluate

the Geneticof Japn. Soc. of Zootech.

The effect of Marker Informationof9 1th annual

meetings

RelationshipSci. : 144.
to evaluate

theof Japn. Soc. of Zootech.

NewEndocrinological

Methodsfor Productionof Germ Cells in Birds

SUSUMU ISM1 , SHUICHI WAKABAYASHI2 and MOTOSHI KDCUCHI1
departmentof Biology,^Department of Biology,

Waseda University,
Nihon University

Tokyo 1 69-50,
Schoolof Dentistry,

Japan
Tokyo 1 0 1 , Japan

Introduction
Despite recent advances in biological techniques, little has been incorporated into
conservation biology.
We have been attempting to apply modern endocrinologicalmethods to conservation biology, especially in helping to breed endangered avian species
includingtechniques for captive breeding programs (Ishii, et ai9- 1995).
To selectindividuals for efficient breeding, from a captive colony of an endangered avian species,
it is necessary to know the gametogenic and endocrine conditions of the ovary and testisof birds in the colony. To estimate these conditions, measurement of plasma levels ofhormones which are directly related to the gonadal conditions is necessary, since direct
observation of the gonad by laparotomy may not be preferred. However, it is also notrecommended to catch such precious birds for collection of blood samples. To solvethese problems, we have developed a method to measure sex steroid hormone levels inavian feces which reflect sex steroid hormone levels in blood. We first present some ofour data showing how accurately fecal sex steroid levels reflect plasma sex steroid levels
in Japanese quail which were used as a model species. We also present an example of
the application of this method to an endangered species, the "Toki" or Japanese crested
ibis {Nipponia nippon).
According to the estimated gonadal condition by fecal sex steroid analysis
method, we can select individuals
for breeding.
If none of the examined individuals
shows an active gonadal condition, we have to stimulate the testis or ovary of someindividuals
in order to promote breeding. For breeding mammals, including manwith
regressed gonads, administration of gonadotropic hormone is commonly used. However,the situation in birds had been different until recently.
Even in domestic hens withregressed ovaries, researchers have not been able to induce ovulation, the final step of theovarian gametogenic activity, by hormone administration.
Accordingly, we developed amethod to induce ovarian growth and ovulation in female birds by administrationof ahormone (Wakabayashi et al , 1 994). The method for hormonal induction of the ovarian
processes - the oocyte development, ovulation and oviposition - is described.

176

Fecal sex steroid analysis in birds
Extraction methods:
Weemployed three different methods, depending on the species, to extract sex
steroids from fecal samples of birds. The simplest method was extraction using water
as the extraction medium followed by radio-immunoassays of the steroids. This method
was used in the Japanese quail. In the Japanese crested ibis, we had to use an organic
solvent for extraction and LH20 column chromatography for removing substances which
disturbed measurement of steroid hormone by radioimmunoassay. A more complicatedmethod was developed to measure sex steroids in feces of the rock ptarmigan (Lagopus
mutus).
Feces of this bird contained much lipid-like
substances which had to be
separated from the steroids before radioimmunoassay.
For this purpose, we used
extraction
with an organic solvent followed by a high performance liquid
chromatograp hy.
Fecal sex steroid study in the Japanese quail:To demonstrate the usefulness of fecal sex steroid analysis method in estimating
gonadal activities,some of our results with the Japanese quail are presented.
In one experiment, we used three groups of male Japanese quail:( 1 ) normal intact adults kept under a long-day photoperiodic
regimen,(2) normal intact adults kept under a short-day photoperiodic
regimen and(3) castrated adults kept under the long-day photoperiodic
regimen.
Meanconcentrations of androgen (testosterone plus dihydrotestosterone)in blood plasma
and feces were compared between the three groups and also between plasma and feces.
The mean concentrations in feces changed in close association with those in plasma(Figure 1). In the next experiment using females, feces were collected from two groupsof Japanese quail hens, one laying at a certain period of the day and the other non-laying.
It is well known, that progesterone in plasma shows a transient rise around the time ofthe ovulation in laying hens. We collected fecal samples from laying hens andprogesterone in the samples was measured. The mean concentration of progesterone infeces had a clear peak around the time of ovulation that was estimated from theoviposition
time in the laying group but no significant
peak in the non-laying group(Figure 2). Thus, we could show that the gonadal activity can be estimated by monitoring
fecal sex steroid hormone. Unlike mammals, birds excrete both urine and feces together.

Figure 1.

Mean concentrations of testosterone in blood plasma (pg/ml) and feces (xlO"1 pg/dropping)innormal adult males kept in a long-day photoperiodic
regimen, normal adult males kept in ashort-day photoperiodic regimen and castrated males kept in the long-day photoperiodic
regimen.

FigureMean concentrations of estradiol-17
beta in blood plasma (xlG~2 pg/g) and feces (pg/dropping)in normal adult females kept in a long-day photoperiodic
regimen, normal adult females kept ina short-day photoperiodic
regimen and immature females.

2.

178

Sex steroids are metabolized to glucronide or sulfate forms before they are excreted.
However, sufficient quantities of the free form (circulating
form ) of the steroids weredetected in feces in our experiments. We suppose that they moved from the circulatory
system into the contents of the digestive duct by diffusion, because they can easily passthrough cell membranes.
Fecal sex steroid study in the Japanese crested ibis:
We applied the fecal sex steroid analysis method to assess the reproductive
condition of a male, Japanese crested ibis. The mean concentrations of androgen forseveral days in three summermonths in 1994 and a period for several days around New
Year's day of 1995 are shown (Figure 4). Compared to the summer months, theconcentration was significantly higher in the period around New Year's day when nuptial
plumage coloration appeared. It is clear that the fecal androgen concentration reflects
well the gonadal activity.
Hormonal induction of the oocyte growth, ovulation and ovipositionin theJapanese quail
Repeated injections of gonadotropin preparations do not induce ovulation in hensof female domestic birds (Wakabayashi
et al, 1994).
We studied changes in theluteinizing
hormone (LH) concentration in blood plasma in Japanese quail. The LH
concentration showed a transient increase and dropped to subnormal levels after several
hours. Then, we decided to use the osmotic pump (ALZET Co., ) for the chronic
treatment of female Japanese quail with gonadotropin.
These birds were kept under ashort-days for a certain period before and during the experiment. Their ovaries wereconsidered to be in the completely regressed condition at the start of the experiment. The
pump was loaded with chicken pituitary
glycoprotein
fraction which is rich ingonadotropins, and implanted into the abdominal cavity. The flow rate of glycoprotein
was 12.5 (ig /hr. This quantity was equivalent to 1.5 \xg chicken LH/hr. High
gonadotropin levels in plasma were maintained over a period of two weeks, when plasma
gonadotropin levels were determined by radioimmunoassay.
The ovary of the females
treated with pituitary glycoprotein
for about two weeks, by means of the osmotic pump,
was well developed and contained a number of oocytes containing the yolk (Figure 5).
However, no ovulation was induced in these females. To mimic the LH surge which

Figure3.

Figure 4,

Change in the mean concentration of progesterone in feces in laying females
quail. The time of ovulation, 1 8:00-20:00,
was estimated from the ovipositionof the Japanesetime.

Comparison offecal testosterone
levels between different seasons in the male Japanese-crested
ibis named "Midori" or "Green". Values are means and their standard errors of two to threesuccessive days in each season.

Figure5.

Ovarian weights of female Japanese quail kept in a short-day regimen, implanted with anosmotic pump loaded with chicken pituitary glycoproteinin the short-day regimen and kept
in a long-day regimen.

induces ovulation in normal hens? weinjected chicken pituitary glycoprotein daily for thelast five days of the osmotic pump treatment in female Japanese quail kept under the
short-day regimen. Four of the 7 treated females laid 7 eggs during the injection period.
One of them laid four eggs and each of the remaining three one egg. All the eggs wereincubated artificially and two of them hatched. The chicks were males and one of themwas confirmed to be fertile by mating with an adult virgin female. We have beensuccessful in stimulating the testis of male Japanese quail kept under a short-day regimen
by this treatment.
Conclusion and Discussion
The fecal sex steroid analysis method is a useful tool to estimate the reproductive
condition of birds non-invasively. Wemayapply this method to monitor the reproductive
condition of birds in feral populations.
Furthermore, in the case of captive breeding of

181

birds, we may select individuals
for breeding from a captive colony of birds using thismethod. However, selection of an appropriate extraction method is important to obtaina satisfactory result.
Our hormone administration
study was the first successful case of hormonal
inductionof a series of ovarian events, i.e. oocyte development, ovulation andoviposition, in birds. Combination of the two different means of hormone administration,
the osmotic pump and injection, is considered to be the cause of our success. However,for the actual applicationof this method for captive breeding of endangered birds,
problems still remain to be solved. First, the success rate of about 40% needs to be
improved. It is also important to clarify the problem of the immunological response tohormones from heterologous species. We are now conducting studies to improve thesuccess rate and also examining the possibilityof using commercial mammalian
gonadotropin
preparations
such as pregnant mare serum gonadotropin and humanchorionic gonadotropin. Preliminary data indicate that these preparations are effective.
References
Ishii,
S., Wada, M., Wakabayashi,S., Sakai.

H., Kubodera,

Endocrinologicalfor artificial

breeding

studiesY., Yamaguchi

N. and Kikuchi,of theibis,

Japaneseendangered avian species in Asia. J. Biosci.
19, 491-502.
Wakabayashi,S., Kikuchi, M., Wada, M., Sakai, H. and Ishii, S. 1992.
ovulationby administrationa short-day

regimen.

Britishof a chicken

gonadotrophin

Poultry

Sci. 33, 847-858.

Induction

preparation

Nipponia

M. 1994.
nippon,anof ovarian growthandto Japanesequail kept underUse of Chimeric
Resources

Animals to Preserve Animal Genetic

AKIRA ONISHI
Departmentof Animal Breeding

Tsukuba Norindanchi

and Genetics,

P.O. Box 5, Tsukuba,

National
Ibaraki

Instituteof Animal Industry,

305, Japan

1. Productionof chimeric animals
Chimeric animals are composite animals containing geneticallydifferent cells
derived frommorethan two fertilized eggs. There are two major techniques for making
animal chimeras : aggregation and injection methods.
1. 1 Aggregation
method8-cell stage embryos are generally used for the aggregation
method. Afterremoving the zona pellucida using 0.5% pronase or acidic Tyrode solution, the embryos
are stuck together with a medium containing phytohemagglutinin.
The aggregatedembryos form blastocysts which are transferred into the uterus of pseudopregnant female
to obtain chimeric offsprings (Figure 1). The aggregation method is simple and efficientfor murine embryos, but it is not suitable for some livestock animals, because of the
difficultyof embryo culture when the zona pellucida is removed.
1.2 Injection method
Blastocyst
stage embryos are widely used for the injection method. The
trophectoderm cells of blastocysts
are broken by immunosurgery using antibodies andcomplement for isolation of inner cell mass (ICM) cells. The isolated ICM cells are
injected
into blastocoelusing a micromanipulator,
and the injected blastocystsaretransferred in the same way as in the aggregation method (Figure 2). Although theinjection method requires skill, a shorter time in embryo-culture and the non-removal ofzona from embryo makes this method appropriate for many livestock animals.
1.3 Production of chimeric pigs
In contrast to the numerous experiments involving chimeric mice, there havebeen relatively few reports of chimeras produced from livestock such as sheep (Fehilly
etal 1984; Butler^al1987), cows (Bran et al 1984; Williams et al 1990; Picard ^al 1990), and pigs (Kashiwazaki
et al 1992).
We examined two injection methods for making chimeras between Chinese

184
Table

1. The production
1994)of piglets

followingR e cip ie n t e m b ry o s

E x p .IE x p . II

blastomere

or ICM cells transplantation

D o n o r c e lls

N o . o f e m b ry o sb re e d *sta g eb re e d *sta g etran sfe rre dMM

4 ‑8 c e ll
8 ‑1
8 6c ecllell

LWML

8 c e l1
8 ‑816c ecl1e l1

17
41 21

M
M
M

b la sto c y stb la sto c y stb la sto c y st

LLL

b la sto c y stb la sto c y stb la sto c y st

13
15
35

*M=Meishan; L=Landrace;(Onishi

et al ,

N o . o f p ig letsb la ckw h itec h im e rall21

LW=Landrace x Large White

Meishan and European Landrace or Landrace x Large White pigs (Table 1). The
Meishan breed has a black coat color whereas the other breeds have a white coat. In thefirst experiment, blastomeres were transplanted into embryos at the 4- 16 cell stage. Of
4 1 transplanted embryos transferred into 3 females, 12 were single-colored,
but no overt
chimeras wereobtained. 2 piglets in 2 litters were derived from injected blastomeres, and
1 0 piglets in 3 litters were derived fromrecipient blastomeres.
In the second experiment,
ICM cells of Day 6 Landrace embryos were injected into the blastocoel of Day 6 Meishanembryos. Of 35 injected embryos transferred into 3 females, 2 overt chimeras of each sex
were obtained in a single litter (Figure 3) (Onishi et al 1994).
2. Analysis of chimerism
The ideal cell markers for analysis of chimeras should be cell-localized,
cell-autonomous, stable, and present in detectable quantities in all cells (McLaren 1976).
In the mouse, the most widely used markers are strain-dependent
enzyme isoforms thatare detectable by electrophoresis.
Although many antigenic and biochemical markers
have been reported in pigs, very few markers are available for characterizationanddistinctionof each breed. Mitochondria are present in all cells, and mitochondria havetheir ownDNA. Mitochondrial DNA (mtDNA) polymorphism in domestic animals has
been reported in many species. In pigs, restriction
fragment length polymorphism of
mtDNA of Chinese breeds is quite different from European breeds (Watanabe et al
1985, 1986; Mikami et al 1988; Takeda et al 1995).
Therefore, we examined theefficiency of mtDNA polymorphism as a cell marker in the analysis of chimerism.

185

Figure1.
Schematic
representationof theprocedure for making aggregation
chimeras.
(a) collectionof 8-cell stage embryos, (b)removing zona pellucida.
(c) aggregation
thepairs, (d) culture to the blastocyst
stage, (e)transfer the aggregatedblastocysts
into theuterus.

Figure2* Schematic
representationofprocedure for making Injection
chimeras,collectionof blastocysts.
(b) Isolation
dlsaggregationof ICM cells, (c) injection
ICM cells Into blastocoel.
(d) transfer
Injected blastocysts
Into uterus.
.

the
(a)
and
the
the

The cloned 3 kb EcoRI fragment of Landrace mtDNA was used as a probe fordetection of Bglll-digested
mtDNA in the chimeric pigs. The Bglll fragment of Meishan(16.5 kb) and Landrace (4.4kb) mtDNA are clearly identified in all organs of the overt
male chimera (Figure 4). The same chimerism was detected in the blood of overt female
chimera. No chimerism was detected in the single-colored
piglets
derived from theexperiment. These results clearly showed that mtDNA polymorphism can be used as acell marker in chimeras (Onishi et al 1994).

Figure3.

Overt chimeras at birth, which were produced by Landrace ICM cells injected
Meishan embryos: (Left) male; (Right)
female. ( Onishi et al. 1994 )Figure4. Detection of Bgffi fragments of mt DNA from various tissues
by southern hybridization.
( Onishi et al 1994 )inin a male overt chimera

3. Use of chimeras; Chimeras derived from ES cells
Chimeras have provided valuable material for studies of embryology anddevelopmental
biology (McLaren 1976).
Recently, chimeras also have played an
important role in genetic modificationof the germ line of animals through use ofembryonic stem (ES) cells (Robertson
1987; Joyner 1993).
ES cells are cultured cells
which are derived directly from the ICMs of blastocysts.
ES cells have normal karyotype
and resemble ICM cells especially in their ability to contribute to all tissues including the

187

gametes in chimera. Great care should be taken in the culture of ES cells for the
maintenance of high differentiation
ability.
The use of feeder layer, together with highqualitymedium, and the accurate time of cell-passage
are important factors formaintenance of ES cells. Following introduction of DNA into ES cells, the targeted ES
cells can be selected and injected into blastocysts
for making chimeras. The targeted ES
cells are transmitted through the germ line of chimeras (Figure 5). Although ES cells area powerful tool for genetic alteration of animals, at present, the establishmentof ES cells
seems to be limited to the mouse. We have not been successful in making chimeras
derived from cultured ICM cells of porcine blastocysts.
Wheeler et al ( 1 994) reported
the live chimeric pigs following injection of long term cultured ICM cells, but they did
not mention the germ line transmission in the chimeras. The study of isolation of ES cells
from livestock animals should be continued.
4. Chimeric pigs for conservationof genetic resources
Obtaining
live offspringfromembryos preserved at -196 °C is
important in animal breeding andconservation of genetic resources.
Recently, live piglets were obtained
from frozen embryos after removing
cytoplasmic lipid (Nagashima et al ,1995).
However, porcine embryos
are still difficult to maintain in longterm storage at low temperatures. In
contrast to the troublesome procedureof embryo freezing, ES cells are easy
to cryopreserve like other tissue
culture cells. In the mouse, theF1(Heterozygote)
productionof offspringfrom
F2 (Homozygote)cryopreserved
ES cells is widely
Figure 5. Diagram to show the production of offspring
carried out through the germ line offrom targeted ES cells, (a) establishmentof ES cells
from ICM cells, (b) introductionof DNA into ES cells,
chimeras. From this point of view, if(c) selectionof targeted ES cells, (d) productionofporcine ES cells are established,itchimeras, (e) breeding to maintain targeted alleles.

188

will be possible
to obtain piglets derived from frozen ES cells by making chimeras.
Nuclear transfer techniques may be the another way to produce complete ES cell-derived
pigs.
References
Brem, G., Tenhumberg,of morulae.
Butler,Theriogenology

J. E., Anderson,

ofovineFehilly,

H., and Krauslish,

H. 1984.

Chimerismin cattle

throughmicrosurgicalaggregation

22: 609-613.

G. B., BonDurant,

R. H., Pashen

chimeras

by inner cell mass transplantation.

C. B., Willadsen

S. M., and Tucker E. M. 1984.

R.

L., and Penedo, M. C. T. 1987.

Production

J. Anim. Sci. 62: 317-324.
Experimental

chimerismin sheep.

J. Reprod.

Fertil.

70:347-351.
Joyner A. L. 1993.
Kashiwazaki,
blastocyst

Gene Targeting.

IRL Press, Oxford,N., Nakao, H., Ohtani,S., and Nakatsuji,

injection

McLaren A. 1 976.
Mikami,method.

Vet. Rec. 130:

Mammalian Chimaeras.

H., Onishi,UK.
N. 1992.

Production

Cambridge

A., Komatsu, M., Otani,

University

T., and Igarashi,

Cambridge,

1988.

Restriction

patternsof mitochondrial

DNA in Chinese

H., Kashiwazaki,N., Ashman, R. J., Grupen, C.G., and Nottle,of porcine

embryos.

Nature 374:

of chimerism

using

pigs

by theJpn. J. Swine Sci. 25: 181-185

UK.
enzyme cleavage(in Japanese).

M. B. 1995.

Cryopreservation

416.

A, Takeda, K., Komatsu, M., Akita,

analysispigs.

Press,S.

Nagashima,
Onishi,of chimeric

186-187.

T., and Kojima,mitochondrial

T. 1994.

deoxyribonucleic

Productionof chimericpigs and theacid as a cell marker.

Biol.

Reprod.

Production

bovine5 1:1069-1075.
Picard,L., Chartrain,

and calves
Robertson,
Takeda,I, King,

W. A., and Betteridge,

by aggregation

E. J. 1 987.
K., Onishi,

Teratocarcinomas

A., Ishida,

and Embryonic

DNA D-loop region

Watanabe, T., Hayashi,Y., Ogasawara,

based

on restriction

Watanabe, T., Hayashi,
mitochondrial

Mol. Reprod.

Stem Cells.

of chimeric

polymorphism.

N., and Tomita,

endonucleasecleavageY., Kimura, J., Yasuda,

DNA: polymorphism,

restriction

embryos

Dev. 27: 295-304.

IRL Press, Oxford,N., Kawakami, K., Komatsu, M., and Inumaru, S. 1995.

pig mitochondrial
pigs

K. J. 1990.

of inner cell masses with morulae.

UK.
SSCP analysisofAnim. Genet. 26: 321 -326.
T. 1985.

Polymorphismof mitochondria

patterns.

Biochem.

Genet. 23: 105-1

Y., Saitou,N., Tomita,T., and Ogasawara,

map orientation,and sequence

data.

of swine embryonicstem cells:

a review.

DNA in13.
N. 1986.

Pig

Biochem.

Genet.

Reprod.

Fertil.

24:385-396.
Wheeler, M. B. 1994.
Dev. 6: 563Williams,

Development

and validation

568.

T. J., Munro, R. K, and Shelton

aggregationof Bos indicus

J. N. 1990.

and Bos taurus demi-embryos.

Production
Reprod.

of interspecies
Fertil.

chimeric

Dev. 2: 385-394.

calvesby

189

Questionsand answers in Session 4

Question to Dr. Ishii.
Q. Dr. Ishii, how do you get individuals
from testicular cells of Japanese Crested Ibis
(Nipponia nipporifl (Naito)
A. Weplan to use nuclei of cryopreserved spermatogonia of a male Nipponia nipponof Japanese origin and exchange nuclei with the nuclei of the Chinese male. Then, we
manyobtain germ cells of the Japanese origin in Chinese individuals.
However,
unfortunately, we have not been able to find germ cells in cryopreserved testicular
cells of the Japanese male which died recently. (Ishii)
Questions to Dr. Takeda.
Q. Dr. Takeda, how many animals are necessary for maintaining the heterozygosityofsmall populations of animals? (Sekikawa)A. This depends on how many generations we are talking about and the level ofheterozygosity, and how manymarkers wehave available. We have to consider eachcase separately. (Takeda)
C. The moreheterozygous the loci, or polymorphic genes are present in a population thelarger the population size is needed for maintaining the genetic structure of the
population. For example the Chillingham
cattle of the United Kingdom has survived
with about 30 females for many centuries because it is highly homogeneous. (Bodo)
Q. Are you using gene-markers to study qualitativeand quantitative
traits? (Astuti)
A. The genotype of the markers are used to estimate the variability of the neighbour lociwhich could be either quantitative or qualitative. However the marker loci should be
qualitative
so that we can distinguish
the genotype clearly. (Takeda)
Questions to Dr. Onishi.
Q. Do you have further plans to make transgenic pigs? (Kagami)
A. At present I have no plans. (Onishi)
Q. Did you observe any heterotic effects in the germline of chimeric pig? (Kagami)
A. Chimeric heterosis for reproductive and growth performance was obvious in the
chimeras derived fromtwo inbred strains of mouse. At present, weplan to examine
relationships
between the degree of chimerism and reproductive traits in chimeric190

pigs. (Onishi)
Q. How did you prove the germline transmission
from your chimeras? Could you
explain how you establishedthe ES cell? (Sekikawa)
A. Meishan has a black coat, while the Landrace has a white coat. In pigs white isdominant to black. Therefore, the progeny test by mating with Meishan bore
establishedthat the white coloured piglets were derived from Landrace oocytes andblack coloured piglets were from Meishan oocytes.
There is one paper about the establishmentof ES cells derived from pigs (porcine)
blastocytes. The authors succeeded in making chimeras using the establishedporcineES cells. However the authors did not mention the transmission of ES cells througha chimera. (Onishi)

TECHNICALREPORTS

Session 5
Use and information

management

Chairpersons
H. Hayakawa

Conservationand Use of the Kacang Goat in Indonesia

MARIA AS TUTI
Facultyof Animal Husbandry,

Jl. Agro Karangmalang,

GadjahYogyakarta

Mada University
5528 1 , Indonesia

Abstract
Kacang goats are the native goats of Indonesia.

Based on the population

females, the present status of this breed is normal from a conservation
to conserved this breed, if any. Conservation

is mostly

perspective.

due to geographic

size of the breeding

Not much have been done

isolation,

historicaland culturalvalue of the breed. Demand of improving

production

and the social changes in the rural life to some extent

influenceresourcesof this breed.

and may threaten

the genetic

necessary to prevent genetic loss and dilution.
breed, the value and potential

This paper discuss

uses along with information

Therefore

a conservation

the reasons and benefits

program isof conserving

this

on performance and management.

Introduction
Three major goat breeds are known in Indonesia, the Kacang goat, the Localgoat, and the Etawah Grade goat. Nearly all goat populations are raised by smallholder
rural farmers. Each farmer raises only a small number.
The largest goat populations are in Java, followed by Sumatra and Sulawesi.
Most of the goat breed populations are dominated by the Local goat which is a crossbred
between Kacang and Etawah Grade goat. The genetic background and development oflocal goat is unknown. People often do not differentiate
the local goat and the Kacanggoat and just call them the Kacang goat. In some places they call the local goat othernames. The local goat can be distinguishedeasily from the pure Kacang goat (Figure 1
and 2)as they show characteristics
between the Kacang and the Etawah Grade goat.
The Kacang goats have low productivity
and crossbreeding
is undertaken to
improve production. However, this breed has not been properly evaluated and fully
exploited, though they support the economy of the smallholder rural farmers especiallyin marginal lands. There is no doubt that this breed plays an important role in theeconomy of farmer households. The value of this breed such as biological,
cultural andscientificuses are many. Some factors have been identified
which threaten the genetic
resource of this breed. There are reasons for and benefits froma conservation program
for the Kacang goat.

194

Goat Populations
and Distribution
More than 90% of the world's nearly half billion goats {Capra hircus) are foundin developing countries. Half in Asia and one third in Africa. These goats are calledmicrogoats.
They weigh less than 35 kg fully grown. They are notable for their highreproductive
rates, rapid growth, early maturity, tasty meat, ease of handling, andtolerance of climatic stress and poor feeds (National Research Council, 1 99 1).
Kacang goat is the one that can be classified
as microgoat. Besides the Kacanggoat in Indonesia there are two other goat breeds, the Local goat and the Etawah Gradegoat. The total population for these three breeds in 1992 was over 12 million head, withan average rate of increase of3.8% per annumfrom 1989 to 1992 (DGLS, 1994).
Goats are distributed
throughout Indonesia, however, they are concentrated
mostly in Java. The distributionof goats in Indonesia is shown (Table 1).
The Local goat is the dominant goat in Indonesia and can be found throughout
Indonesia.
This breed is a crossbreed between the Kacang goat and the Etawah Gradegoat. The genetic background of the crossbreed is unknown and varies due to uncontrolled mating.
The distributionof purebreed Kacang goat is limited to certain areas especiallyin the marginal lands and in the isolated hilly and mountainous areas without easy roadaccess.
Separate population data for each goat breed is not available yet, but it isestimated that in 1992 the purebred Kacang goats numbered approximately 2 million
head, and the Etawah Grade goat about 0.5 million head, and about 9 million Local goats.
To obtain an exact number of each breed is not easy because many reports do notdifferentiate
the Local goat from the Kacang goat. In some places the Local goat has alocal name such as Jawa Randu in Central Java, Bligon in Yogyakarta region, Krikil in
Sragen, Marica in South Sulawesi, and Gembrong in Bali.
In Central Java the population of Kacang goats is concentrated in the north of
Bengawan Solo river, on the mountain range of Kendeng and Kapur Utara in the districtof Sragen, Grobogan, Purwodadi, Blora and Rembang. Other areas can be mentioned
like Lebak and Serang districts in West Java, district of Jambi, Palembang, and Lampung
in Sumatra.

195

Table 1. Distributionof goats in Indonesia

in 1989

to 1992 (million

19 8 919 90

19 9 1

19 92

S u m a t ra

1 .1 2 4

2 .2 5 6

2 .3 5 3

2 .4 7 0

Java

6 .5 4 1

6 .6 6 9

6 .6 9 5

7 .0 3 5

K a lim a n ta n

0 .1 6 7

0 .1 7 1

0 .1 7 8

0 .1 9 5I s la n dhead)

S u law e si

1 .0 2 8

1 .0 7 5

1 .1 0 6

1 .1 1 0

W e st N u s a T e n g g a ra

0 .3 8 0

0 .3 6 6

0 .3 8 6

0 .3 5 9E a st N u sa T e n g g a ra

0 .4 5 7

0 .4 4 7

0 .4 3 4

0 .5 2 8M a lu k u a n d I r ia n J a v a0 .2 0 6

0 .2 1 4

0 .2 3 0

0 .2 4 6

E a s t T im o r

0 .0 9 1

0 .0 9 7

0 .1 0 0

0 .1 1 6I n d o n e s ia

1 0 ,9 9 4

1 1 .2 9 5

1 1 .4 8 3

1 2 .0 5 9

Source:

DGLS,

1994.

Kacang Goat Native Breed to Indonesia
Kacang goat is a native breed to Indonesia. Devendra ( 1 983) reported that thisbreed are also found in Malaysia and Philippines.
Before the 1920fs goat's in Indonesia
were the Kacang goat. In 1923 , during Dutch colonization, the Etawah goat (Jamnapari)
was introduced from India (Soepardjo,
1 980).
Etawah goats have a larger body frame, long hanging ears, convex face, large
horns, the udder are well developed and they are used for milk prodution. The Etawah
breed was imported with the intention of improving the meat and milk production of thenative Kacang goat. These imported goats were then bred and developed in the breeding
center of Pangarasan (Madura), and from this center the Etawah goats were distributed
to other animal husbandry centres such as Cirebon, Pekalongan, Purworedjo, Blitar,
Nganjuk, Nusa Tenggara, North and South Sulawesi, and North Sumatra (Soedjai,
1 952
and Aldjufii, 1971). From that time the Etawah grade goats were known as descendants
from crosses between the native Kacang goats and the Etawah bucks. Then during thedevelopment of the goat population, Etawah grade goats were used for crossing with
Kacang goats. Local goats are the results of uncontrolled crossing with Etawah gradegoats. Due to the early history of the distributionof Etawah goats, in the regions where
Etawah goats were distributed
and in the surrounding regions nowonly Local goats canbe found. The population of Local goats has increased dramaticallyand is widelydistributed
because of the government support for crossing the Kacang goats with the
Etawah grade goats.
Kacang goats are well adapted to the agroclimaticconditions of Indonesia andany selection that has taken place over the years has been entirely by natural selection.

Table 2.

Production

and reproduction

performance of Kacang goat

Crossing the Kacang goat with other breeds might cause a dilution and loss of the genes
related to adaptation and other valuable genes as yet unidentified.
Purebred Kacang goat has a certain characteristics
which is manifested in thesmall and compact body frame, erect ears, flat nose, short neck, short horns, and the backrises slightly higher than the shoulders. Mature males and females weigh approximately
25 kg and 20 kg respectively with the average height at wither of 60 - 65 cm in males and
56 cm in females (Devendra and Burns, 1970). Kacang goats are known as a prolificgoat and they produce kids all year around (Devendra, 1983).
Management and Performance
Goats are raised by smallholder

rural farmers. One in five farmers in Indonesia

Figure 1. Typical Kacang goats in the back in theforeground goats with white faces show theinfluence of crossing which is also indicated by
the change in the ear type.

Figure 2. Kacang goats at livestock
Jumberlawang district of Sragen.

marketraises goats or sheep (Soedjana,1993).
The herd size is usually 2 to 7 head and 2 to 3
head is most common.Farmers raise the goats under traditional
management systems,
and goat rearing is only a secondary farming activity. Goats are reared under grazing andbrowsing systems, cut and carry systems and farmers in urban areas rear goats underscavenging systems. Goats are usually grazing and browsing along the road sides, banks,
wastelands and uncultivated rice fields in the morning and penned at night. Under cut
and carry system farmers give cut-grass, legume tree and other tree leaves and sometimes
the goats graze. In urban areas goats scavenge in the market, along the road and in
garbage dumps. At night goats are tethered in the barn or the farmer's house. Only few
farmers use a raised floor pen.
Not all farmers owntheir ownbuck. There is uncontrolled mating, goats are free
to mate indiscriminantly.
Generally does nurse their kids until weaning which is after
three months.
Little has been done in disease prevention and control. Loses of kids are morecommonthan loses of adults. The most commondiseases are scabies, diarrhoea, pinkeye, orf, bloat and gastro intestinal
parasite investation.
Rural farmers use traditional
medicine to cure disease. The use of traditional
medicine varies from region to region.
Near cities, farmers use traditional
medicine less frequently due to the availabilityofveterinary services that help farmers by monitoring diseases (Sutama and Djajanegara,1992). In traditional
medicine, diarrhoea is treated by giving bamboo leaves or ground198

guava leaves mixed with boiled water. Bloat is cured with ground ginger rhizome mixed
with black coffee and boiled water. To overcome the internal parasites farmers use
ground black tumeric (Temum hitam) rhizome mixed with water for drenching. Scabies
usually is treated by rubbing with a solution of lubrication
oil and sulphur.
The performance of Kacang goats has been summarized in Table 2. Data in
Table 2 indicates that the productivityof Kacang goat is low in terms of weight and gain.
The Reasons and Benefits of Conservation
Bodo (1990)
suggested two reasons why a population should be conserved,
these are:
1. The endangered status.
2. Genetic value.
Further, Diwyanto and Handayani ( 1 995) stated that the conservation of native livestockin Indonesia could be justified
for economic, biologicaland socio-cultural
reasons. All
these reasons can be used to justify the conservation of Kacang goats. Kacang goats area valuable genetic resources especially in relation to genetics of adaptation.
Kacang goats
are adapted to the local climatic conditions, the poor quality of feed resources, themanagement system under which they are kept, and resistance to some infectious andparasitic diseases.
The status of populations,according to Bodo (1990), can be categorized into
five groups based on the number of the breeding females i.e. normal, insecure, vulnerable,
endangered, and critical. The present status of Kacang goat population can be categorized
as normal.
Although the status is normal this breed as a purebred is influenced andthreatened by several factors. These factors are:
1. Improving productivityof crossbreeds.
This has been implemented through crossbreeding
Kacang with Etawah grade
bucks, and has been carried out over decades with the objectivesof increasing the size ofthe animal and improving milk productivity.
Crossbreedingwith uncontrolled mating
will cause a dilution and loss of the genes related to adaptation and other genes that mightby valuable in the future. The Kacang goats cannot maintain their original type, size,
conformation and performance due to crossbreeding.
Efforts should be made to improve

199

the production of Kacang goats while still maintaining the purity of the breed. On theother hand, crossbreeding with other breeds can be used as a tool to produce commercial
goats to meet the market demand.
2. Socio-economic changes in the rural areas.
More people especially young people are leaving rural areas, looking for a better
life and job in the nearby cities. Farmer's interests have changed due to modernizationand they are becoming reluctant to raise goats.
Rapid changes in transportation
infrastructure
make it easier to purchase and bring other goat breeds to rural areas.
3. Animal health problem.
Disease and parasite control are not commonly practiced. Mortality is quite highin young kids. Though the Kacang goats are believed to be resistance to disease andparasites long exposure to disease and parasite will cause mortality.
The benefit of conservation can be viewed from 1. The economic uses, 2. The
biologicaluses, 3. The cultural uses, and 4. The scientific uses.
1. The economic uses
The economic uses of a given stock are determined by the performance, theadaptation, and disease resistance (FAO, 1 990). The Kacang goats are believed to havea potential for adaptation and disease resistance.
The performance of Kacang goats interm of production is low but this goat has some economic advantages for the smallholder
rural farmers. Like other microbreeds they are less expensive to buy, less of a financial
risk to maintain, give a faster return for investment, use space efficiently,
are easy andcheaper to manage(National Research Council, 199 1). These goats support the economyof the farmers. In Java, goats contribute 10 to 30% of the farmers income (ISRN, 1992).
Further goats are used to buffer investment risk and provide capital and cash. To attaingreater economic benefits farmers should be informed of how to be good at animalhusbandry including the raising of goats. Further, the advantages of the genetics ofadaptation and disease resistance should be exploited.
2. The biologicaluses
Kacang goats as the native breed of Indonesia,

enrich the biologicaldiversity

of

200

the country. Kacang goat might have some valuable genes related to some biologicaltraits that have not been used by farmers such as the gene for prolificacy.
Somebiological traits might not be found in exotic animals and they will be needed to develop
newcrossbred animals.
Kacang goats are particularly
useful for meat production.
Other uses are for
producing organic fertilizers(manure) and for producing hides. As a source of animal
protein, goat meat is an appropriate source of animal protein to support the national
program of improving per capita protein consumption in rural areas. It is a relatively
inexpensive
meat that rural households
can afford and there is increasing
meat
consumption as household expenditure increase (Soedjana,
1 993).
At present the national goat meat production is 76,590 tons, which is 1 6% of thetotal meat supply from ruminant livestocks (DGLS, 1994).
3. The cultural uses
Kacang goat are used in religious ceremonies. People sacrifices the Kacang goatsat funerals, to celebrate the the birth of a child, and as a brideprice in marriage. All ofthese are rooted to the Indonesian cultural heritage especially important in rural life.
4. The scientificuses
Kacang goats are used as experimental animals in production research, genetic
research and nutrition research. Recently, researchers are using fistulatedgoats to study
in sacco and in vitro digestibilities.
Compared to fistulated large ruminants the use offistulated goat is cheaper and needs less space, so goats are becoming more popular forsuch experiments.
In the future with the applicationof biotechnology
the genetic
potential can be exploited for the benefit of mankind.
Conservation of Kacang Goat
Livestockconservation must be conductedwithin the livestocks
own
environment. A sufficient population size for each genotype is necessary. It's known
that the preserved genotype will always be the result of combined effects of genetical andenvironmental variables, the latter being peculiar
to the breeding environment. The
breeding environment where the livestock has evolved or environments that are thought
to be similar to the original one will enable them to live and reproduce (Matassino et al.
1993).
Oneway to conserve the genetic resources of livestock is by conservation of

102

individuals

either in situ or ex situ.
In situ conservation or on-farm conservation might be the appropriate
way for
Kacang goats to be conserved whereby populations are raised by the farmers in their own
environment.
The most effective on-farm conservation is that which is nationally promoted
and combines elements of existing national programs(Diwyantoand Handayani, 1 995).
A government-sponsored plant and animal conservation effort exist in Indonesia such as
the National Genetic Resource Committee (Komisi National Plasma Nutfah/KNPN).
The national program should pursue the genetic improvement of Kacang goats while on
the other hand maintaining the genetic diversity of the breed. KNPN as a single entity
should initiate the efforts and give suggestion and recommendations concerning thebreeding policy of government.
At the local level where the pure Kacang goats are found, a group of farmers
with the same interest should be organized. Local government help to such groups withknowledge of animal husbandry practices and means to prevent the use of other goat
breeds for breeding. Commonly farmers buy and sell their goats in the nearby livestock
markets which are located within the radius 10 to 15 km from their home. In the
livestock market farmers purchase the bucks or does. To prevent crossing with otherbreeds, the local livestock services should establisheda buffer zone and make a rule that
only Kacang goats are allowed to be sold inside this buffer zone.
Implementation of on-farm conservation includes the national policy to conserve
the Kacang goats, evaluation at the phenotypic and genetic level to know more about
biologicalcharacteristicsof the Kacang goats, to establish an organization for on-farmconservation, and training of technical personal to maintain the program.
Conclusions
The population of the Kacang goat is decreasing due to crossing with Etawah
grade buck with the intention of improving productivity.
Kacang goats have a genetic
value for adaptation and prolificacy. Continuous and uncontrolled crossing will cause theloss and dilution of these valuable genetic resources.
These reasons and the benefits justify conserving Kacang goats. Oneway toconserve this breed is by on-farm conservation. A breeding policy for the Kacang goatshould be establishedto support the efforts of conservation.

202

The genetic potential of Kacang goat should be properlyin the existing conditions where they are exposed.

evaluatedand exploited

Seminar Penelitiandan Pendidikan

References
Aldjufri,S. 1971.

Susu Kambing

Sebagai

Sumber Protein

Keluarga.

Peternakan Ciawi, Bogor, Indonesia (Indonesian).
Astuti, M., Sunardi, Arinto. 1994. The Impact of Outbreeding
Returns
Astuti,in Kacang

Goat.

Research(Indonesian).
M. 1995. Growth and Survivability
Animal Science. Special
University,
Yogyakarta,

Bodo, I. 1990.

Methods

Resources:

and Crossbreeding

report.

Agriculturalof Preweaning

Edition a Publication
Indonesia.

and Experienceswith in situ Preservationin the Tropics.

Farnham Royal, Bucks, England.
Devendra, C. 1 983. Goats: Husbandry and Potential

in Malaysia

Book on Livestock.

Diwyanto, K. and S.W. Handayani.

Southeast

Asia

at the Third

Indonesia,

22-24 August 1995.

Manual on Establishment

Organization
ISRN.

Strategicpresented
FAO. 1990.
1992.

Directorate

1 995.

of Farm Animals.

Project

In Animal

Ministryof Agriculture

on Geneticof Animal

Services,

Conservation

of
Mada

Genetic

Anim. Prod, and HealthGeneral of Livestock

Symposium

GadjahCommonwealth Agricultural

On-Farm Livestockand Operation

Rome, Italy,of Animal Husbandry,

Development.

No. 80. FAO, Rome, pp 85-102.
Devendra, C. and M. Burns. 1970. Goat Production

DGLS. 1 994. Statisticaland Economic

Management

Kacang and Kacang Cross Kids. Bulletinof the Faculty

A Global Programme for Suntainable

on Biologic

Research

Paper

Bureaux

Malaysia.
Jakarta,Indonesia.

in Indonesia.

Paper

Serpong,

Jakarta,Resources.

Genes Banks. Food and Agriculture

p. 67.

Summary and Conclusionsof the Workshop

on the Potential

and Developmentof Goats inEastern Indonesia. Indonesia Small Ruminant Network Newsletter. Bogor, Indonesia, vol. 3 :7-8.
Matassindo, D., A. Cappuccio, F. Grasso and M. Palazzo. 1993. Conservationof Animal Germplasma at
Risk ofExtination
National

Soedjana,
Soepardjo,

The centre for the Defense of Animal Genetic Resources

In Animal Genetic Resources Information. FAO, Rome, pp 27-48.
Research Council. 1 99 1. Microlivestock:
LittleKnown
Small Animals
Future.

Soedjai,in Italy:

National

R.A. 1952.

Academy Press, Washington,

Ilmu Hewan Ternak di Indonesia.

T.D. 1993.

Small

Indonesia

Small Ruminant

T. 1980.

Kambing

Pertemuan Ilmiah
Sutama, IK. and A. Djajanegara,

Ruminant

Pergjzi

G. Kolff&

Bogor, Indonesia.

Sumber Penyediaan

Pangan. Yogyakarta,

1 992. Traditional

Co. Bandung (Indonesian).

in Java, and Effect on Income on Consumption.

Network Newsletter,

Indonesia.

Medicine:

Systems Network for Asia. Bogor, Indonesia.

Economic

D.C.

Meat Production

Perah Sebagaiwith a Promisingof Circello.

26 Januari

A Feed AdditiveVol. 3:3.

Vol. 4: 5-7.

Air Susu di Pedesaan.
1980
Small

Makalahpada(Indonesian).
Ruminant Production

Survey and Developmentof Experimental

Animals

KAT SUHIRO FUKUTA
Department of Bio Defense Research Division, National Instituteof Animal HealthKannondai 3-1-1 , Tsukuba, Ibaraki 305, Japan
Present address:
School of Agricultural
Sciences, Nagoya University
Furo-cho, Chikusa-ku, Nagoya 464-0 1 , Japan

Introduction
Animal experiments occupy an important position in the broad field of life
science.
In experiments we can use any animals, but they can be classified
into 3categories, that is, laboratory animals, domestic animals and captive wild animals. The
laboratory animals are developed as sophisticatedtools for use in research. They havebeen bred under genetic, microbiologicaland environmental control. Domestic animals
are easy to use in experiments, but most of them have been bred without considering their
genetic or microbiologicalbackground. The term "experimental animals" is used in a
broad sense for animals used in experiments, including laboratory animals and animals
in the process of development for laboratory use. Generally experimental animals havebeen developed for use in the field of medical and pharmaceuticalscience as a substitutefor human being. On the other hand, in the field of veterinary medicine and animal
science, domestic animals are targets for study and can be used for experiments.
However, large domestic animals are disadvantageous for use in the laboratory.
In thepresent paper the survey and development of experimental animals for veterinary
medicine and animal science of use as substitutes for large domestic animals is described.
Worldwide, there are about 4,500 species of mammals, 8,600 species of birds,
6,000 species of reptile, and 3,000 species of amphibia.
However, there are only 10
commonly used experimental animals. The most popular experimental animals arelaboratory rodents like mice and rats. They are geneticallyand microbiologicallysophisticated.
There are more than 400 inbred strains of mice (Festing,1993).
Microbiologicallycontrolled rodents, such as germ free and SPF (specific pathogen free)mice, have been established.
Therefore laboratory mice have been called "living test
tubes", and they have contributed very much to progress in life science.
The advantage of using domestic animals in veterinary medicine and animal
science research, is that the results of experiments can be directly applied to the field.
While, the disadvantages of domestic animals for use in research is their large body size
which affects :-1. Handling and management requires experts;
2. These animals are costly to purchase and maintain;
3. For chemical and irradiation experiments large doses are required which is dangerous
for experimenters ;4. Facilities
and equipment have to be prepared especially for each animal.
Small laboratory rodents have been used even in the field of veterinary medicine
and animal science as well as domestic animals. However, they are omnivorous animals
having a single stomach. Their physiologicalcharacteristics
are quite different fromthose of domestic ruminants. For these reasons, new experimental animals to act as
substitutes for domestic ruminants are needed in the field of veterinary and animal science(Goto, 1986a).
Survey and development of experimental
animals
As candidates
for experimental
animals as substitutes for domestic animals,
particularly
for ruminants, we can classify
them to 4 categories>
1. Small herbivorous rodents with a complex
stomach harboring cellulytic
bacteria;
2. Small- or microbreeds of livestock;
3. Small wild relativesof the domestic
ruminants ;4. Transgenic
mice
added
with
characteristics
of ruminants introduced;
In the following discussion the first three
approaches will be discussed.
The steps in the developmentofexperimental animals are shown (Figure 1).
First of all, we develop a list of as many
animal species as possible as candidates to be
Figure1.
Process of developmentofdeveloped, from bibliographicalsources. In
experimental animal. *These steps can be
the next step, the listed animals are examined
omitted for indigenous animals.
in detail by bibliographicalsurvey and byTable1. List of microlivestock

Table 2. Small wild ruminants

206

Table 3. Body size of small wild ruminants

P u dup̲ u d .u p u d ,uH lee na gd th& (cb om d)yh Se igh oh ut ld(cemr )B^ o d,y w e .g
. h, t, (kn g ).

80 ‑9 0

38

7 ‑9

^

4 5 ‑ 50

4 ‑6B lu e d u ik e r c ep h a lo p h u s m o n tic o la
K irk 's d ik d ik M a a o q u a k ir k ii

55

65

37 . 4 5

3 .4 5R o y a l a n te lo pN e e o tra g u s p y g m a e u s

4 5 ̲ 55

2 0 ̲ 2g

1 5 ̲2C h in e se m u n Mtjaucn tia c u s re e v e si

65

4 Q ̲ 5Q

15

T a b le 4 . B o d y siz e a n d w e ig h t o f m o u se ‑d e e r (F a m ily ; T ra g u lid a e )H iee na gd th^ & (cm,b o d)/yh, eS ig
.h oh, utw ld(cemr ),B^ o d,yJ w e ig
.6 hu t, (kvn g& )J,

A fri c a n w a te rH cy heme v oros ta
c hinu s a q u a tic u s

‑.

‑

3S

40

7

"M

In d ia n c h e v roTtar aing u lu s m e m in n a

45

55

25

30

2 2

2 ?

L a rg e r m o u se‑d
T r ae ge ru lu s n ap u

5Q ̲7 5

30

35

3

5

L e s se r m o u seT‑dr ae geru lu s m v a n ic u s

30

2Q

25

0 ?

47

2

T a b le 5 . R e p ro d u c tio n tra its o f le sse r m o u se ‑d e e r
B re e d in g se a so n
E stro u s c y c le
M a itin g p e rio d
T h e e a rlie st ag e o f c o p u la tio n (m a le )T h e e a rlie st ag e o f c o p u la tio n (fe m a le )T h e sh o rte st g e sta tio n p e rio dA c tu a l g e statio n p e rio d
P o st‑p a rtu m e stro u s
L itte r siz e

N o , c o n tin u o u s b re e d e r
14 ‑1 6 d a y s
2 day s166 day s
2 58 day s
132 day s
134 ｱ 2 day s
Y e s , 3 0 m in u te s a fte r p a rtu ritio no n e , ra re ly tw ofield survey in their actual habitats, as occasion demands. Judging the probability
andvalue to develop the animals, we make a decision on a few animal species as candidates
as experimental animals. In the case of overseas animals, it is necessary to check thestatus of the animals in their native countries, because the animals of interest to be
developed are often rare and protected by law. For introduced animals export and importpermission
is necessary. Obtaining export permission of animals is often the mostdifficultstep in the development of animals. Recently, some governments have atendency to refuse export of animals to foreign countries, whether the animals are rare

207

and protected or not.
Import permission is also difficult
to obtain.
For even-towed ungulates orartiodactyls,
Japan rarely gives import permission, since protection of livestock fromserious infectious diseases is a priority.
Foot-and-mouth disease is most feared due toits high infectivity.
Consequently, animals from Southeast Asia, Africa and SouthAmerica are prohibitedfrom import into Japan in principle.
However, in the case ofacademic research or display at zoos, import permission may be issued after rigorous
quarantine inspection.
Before applying for import permission,
we must gather
information about the past and present hygienic state in relation to animals of theexporting country, then obtain an official certificate of the animals health. With both theexport and import permission, animals can be transferred to Japan, and they are inspected
at a quarantine station for 14 or 35 days depending on animal species and hygienic stateof the exporting country. After that they can be introduced into facilities
for the animals
maintenance. During the first 3 months after the introduction,
if the animals health alters
the quarantine office should be informed.
On rearing of the introduced animals, we must improve food, cage and others to
become moreadequate for them. We should not only rear the animals successfully, butalso proliferatethem effectively.
For the successful breeding, we studied theirreproductive
traits and behavior.
Pairing for reproduction is very difficult to wild
animals, because they are often solitary and struggle each other at the time of pairing.
Some species permit mating partner at only the breeding season in the wild.
While establishinga breeding colony, the animals must be tamed to enableeasily handling and their physiologicalcharacteristics
must be studied. Finally we should
distribute the animals for use in experiments.
Experimental animal candidates( 1) Small herbivorous rodents with complex stomach
Small rodents are useful animals for experiments, because of their small bodysize, short life span, early sexual maturity and large litter size. They are very convenient
for researchers in the field of veterinary medicine and animal science, if small rodents
with complex stomach similar to rumen are developed as replacement for ruminants. The
vole, Microtus, is a herbivorous rodent having a stomach which consists of two
compartments, forestomach and pyloric stomach. The forestomach is covered with

208

stratified epitheliumand cellulolytic
bacteria are harbored there (Kudo & Oki, 1984).
More than 50 species of Microtus spp. distributed
worldwide. Among them, Microtus
oeconomus, M. montebelli, M. arvaris, M. agrestis, M. ochrogaster and M
pensylvanicus have been bred in laboratories.
In NIAH we have maintained Japanesefield vole,Microtus montebelli (Figire 2), for more than 1 5 years (Goto, 1986b).
Duringtheir maintenance, some mutations occur in the colony and are maintained as mutant
strains.
They are a good model substituting
for large domestic ruminants to use innutritional
physiology(Kudo & Oki, 1984). However, the limitation of the vole is that
it is susceptible
to domestic animal diseases.
(2)

Microbreeds of livestock
There are many kinds of small livestock in the world (National Research Council,
1991). Most of them have been maintained as native breeds in isolated rural areas. Withadvances in communication and trade, these traditional
breeds are declining.
A few
microbreeds have been developed by selective breeding.
Table 1 shows a list of
microbreeds of cattle, sheep and goat Amongmicrocattle, Mini-Brahman (Bonsai cattle)
is the smallest breed which has been developed in Mexico by selective breeding byProfessor Berruecos and his colleagues.
However this breed is more than 130kg andinconvenient for experiments.
Small sheep and goat are about 15kg in adult body weight. Actually the West
African dwarf goat and Shiba goat were bred for use in experiments. In Japan the Shibagoat has been examined for many physiologicaltraits.
Katjang is a small goat
distributing
through Southeast Asia, China and Pacific islands.
The Katjang goat ofIndonesia (Figure 3) is also a candidate for an experimental animal, because of their small
body size, easy handling and good reproductive
performance (Fukuta, 1993).
Theadvantage of the microgoat as an experimental animal is that maintenance and handlingis already established.
The size is adequate for training medical students surgery. In
future microgoat and microsheep will prevail as experimental animals in veterinary
medicine, animal science and also in medicine.
(3)

Small wild ruminants
Many kinds of small ruminants, order Artiodactyls,
suborder Ruminantia, aredistributed
through different families and genera (Table 2). These animals inhabit Africa,

209

South America and Southeast Asia. Table 3 and 4 shows their body size and weight.
The smallest is lesser mouse-deer of Southeast Asia and royal antelope of Africa. We
have little information about the royal antelope. The mouse-deer consists of one familyof 2 genera and 4 species. Mouse-deer inhabit Asia except one species, the African water
chevrotain (Dubost, 1 986).
Amongthe small wild ruminants, blue duikers have been bred at the Pennsylvania
State University (Goto, 1986a). The lesser mouse-deer is maintained at the Institute for
Medical Research (IMR), Malaysia; Universiti Pertanian Malaysia (UPM), Malaysia; andNational Institute of Animal Health (NIAH), Japan (Fukuta, 1 996).
Breedingof lesser mouse-deer at NIAH
Lesser mouse-deer, Tragulusjavanicus,
is the smallest ruminant in the world.
Adult body weight is about 1.5 kg and rarely over 2 kg. The lesser mouse-deer has
neither horn nor antler (Figure 4). The male has long upper canines as tusks. We areinterested in this animal and plan to develop it as an experimental animal as a substitutefor domestic ruminants. Fortunately we have been able to import lesser mouse-deer
twice from Malaysia. The first time 2 pairs were introduced from IMR in 1989, andsecond time 15 heads were introduced from UPM in 1995.
To maintain lesser mouse-deer, weuse stainless steel rabbit cages as do IMR and
UPM. The cages are two cages (57x57x47cm) joined together, and a partition is able
to separate them into 2 compartments (Figure 5). At UPM half of the connected-cages
are used to keep a pair and good reproductive performance was achieved(Kudo et ah,
1996). The animals are fed on rabbit pellet feed, banana, carrots, long beans and broken
cubed hay for the first few years, then the array of food was reduced to rabbit pellet feed,banana and cubed hay. Nowadays wegive rabbit pellet feed, sweet potatoes and broken
cubed hay and supplemented with carrots.
The lesser mouse-deer rests in the daytime and moves actively around sunrise andsunset, but they are not nocturnal which is described in many books (Fukuta, 1996). The
stomach is large and occupies most of the abdominal cavity.
It consistsof 3compartments; rumen, reticulum and abomasum (Figure 6 and 7). A small vestigial
omasumis recognized between the reticulum and the abomasum. The reticulo-rumen
volume is larger than that of goat and cattle. Protozoa and large bacteria are found in therumen. Cellulolysis
ability of the rumen bacteria in the lesser mouse-deer is highFigure2. A Japanesemontebelli.

field

vole,

Microtus

Figure 3. Katjang

Figure 4. Lesser mouse-deer, Tragulusjavanicus,
maintainedat NIAH. Male (front right) has
long upper canine tooth acts like a tusk.

goat in Indonesia

Figure 5. Stainlesssteel cages for lesser
mouse-deer. Two original rabbit cages areconnected.

FigureFigure 7. Schematic drawing
lesser mouse- deer.

Figure 8. Young lesser mouse-deer of 2
months old. The animal is tamed by handfeeding.

of stomach of

6. Rumen of lesser mouse-deer.

212

compared with those of cattle and buffalo (Kudo et al , 1996). The reproductive traitsof the lesser mouse-deer are summarized (Table 5).
The disadvantage
of lesser mouse-deer as an experimental animal is its longgestation period and only one individual per litter. However, they can become pregnant
continuously by post-partum estrous and deliver young twice or three times a year.
Therefore, the animals proliferate
more effectively than large domestic animals, if abreeding colony is established.
Although the lesser mouse-deer is said to be delicate andshy, we can tame them by hand feeding (Figure 7).
Conservation of animal species and the development of experimental
animals
Animal welfare groups target animal experiments. For these groups thedevelopment of experimental animals might be considered exhaustion of animal
resources. To develop new experimental animals, however, it is necessary to establish
methods to maintain animals safely and in conditions where they can proliferate
effectively under human control. Rare animals in the wild, are threatened with extinction.
Experimental animal research can give insights into how best to proliferaterare species.
In the past, the procedure of rearing and breeding experimental animals has helped
rescued rare animals from extinction.
For example, in the case of the volcano rabbit(Romerolagus diazi) which inhabits restricted areas of mountains around Mexico city
they were threatened with extinction.
Several pairs of the rabbits were introduced into
Japan and proliferated by researchers who major in experimental animals (Matsuzaki etal, 1982). Young volcano rabbits were exported back to Mexico from Japan. For rareanimals in the wild, the procedure of rearing and breeding of experimental animals may
contribute to their conservation.
Acknowledgments
The author wish thank Dr. N. Goto, former Professor of Animal Breeding,Dr. H. Kudo, National Instituteof Animal Husbandry, for their helpful suggestions
Lesser mouse-deer were obtained
S. Jalaludin,

vice-chancellor,from Dr. N. H. Fuzina, Institute

Universiti

Pertanian

Malaysia.

for Medical

Kobe University, andand encouragement.

Research,

Their help is gratefully

Malaysia,

and Dr.

acknowledged.

References
Dubost,G. (1986).

Tragulidae.

In: The Encyclopedia(ed.),

Vol. 4, Heibonsha

Ltd. Publ.,

Festing,

M. F. (1993).

Internationalof Animals(Japanese

Edition),

MacDonald,

D. W.

MRC Toxicology

Unit,

Tokyo. Pp. 72-73.

Index of Laboratory

Animals,

6th edition,

213
University
Fukuta,K. (1993).
laboratoryof Leicester,

Leicester,U.K.

Investigationof animal

geneticof Agrobiological
Fukuta,resources

in Indonesia

for the developmentof new

Survey Report for Animal Genetic

Resources,

No. 4, 1 -4 1 , National

Institute

Resources,

with English

summary).

animals.

K. (1996).

ReportTsukubaon the breeding(in Japanese

statusof mouse-deer,

introductionof lesser mouse-deer, Tragulusjavanicus,
Genetic Resources, No. 6, National Instituteof Agrobiologicalin Japanese
Goto, N. (1 986).

with English

Candidates

Herbivora,

animals

ruminant.

Experimental

specieslaboratory

animals:

Reproduction;and nutritional

physiology.

Veterinaryas new herbivorous
tract;

K. & Jalaludin,

S. (1996).

The smallest

ruminant,

javanicus).

In: Tropical

Ruminant

and Microbiology(in press).

diazi)
National

M. & Kamiya, M. (1982).

in captivity.

Experimental

research council (1 99 1 ). Microlivestock:
Future.

National

Herbivore,

and its characteristics.

Kudo, H., Fukuta,T., Saito,

Experimental(in Japanese).
Microtus

bacterial
flora and fermentationin the digestive
Research Communications.
8: 77-91.

Matsuzaki,

and theto Japan. Survey Report for Animal
Resources, Tsukuba (in press, andanimal for domestic

field vole as an experimental

1 1 : 63-74

Kudo, H. & Oki, Y. (1984).

spp., in Malaysia

summary).

of the small laboratory

1 1: 9-14 (in Japanese).
Goto, N. (1986).
The Japanese

Tragulus

Breeding
Animals,

and rearing

lessermouse deer {Tragulusof the volcano rabbit(Romerolagus

31 : 1 85-1 88.

Little-known

Academy Press, Washington,

Small Animals
D.C.

with a Promising

Economic

Information System for Animal Genetic Resourcesin the Asian Region
YUTAKA YAMADA1and DAVID E. STEANE2
department of Animal Production,
Hokkaido National

Agricultural

Experiment

Station

Hitsugigaoka
1 , Toyohira-ku, Sapporo, Hokkaido 062, Japan
2GCP/RAS/144/JPN,
FAO Regional Office for Asia and the Pacific
39 Phra Atit Road, Bangkok, Thailand
Summary
The Food and Agriculture

Organizationof the United Nations(FAO)

recentlydevelopeda Special

Action Programme for Global Management of Animal Genetic Resources.

The establishment

Animal Diversity

components of the Programme.

Information

System (DAD-IS)

is one of very importantof Domestic

In addition,
the FAO has started a regional project, supported by the Japanese government, entitled"Conservation and Use of Animal Genetic Resources in Asia and the Pacific". This project covers 12 Asian
countries and is for 4 years. One of the major activities
is to make an inventory of domestic animals and toobtain better population
data in the region. The Regional Databank will be establishedat the project officein Bangkok.

Furthermore,coordinating

institutions

an Asian Network will be established.

in each of the twelve participating

centers, the project office in Bangkok is expected
DAD-IS (Global

Center)E-mail system will be set up at specificcountries.

to be regional

These institutions

will act as national

center and this Network will be linked

to thein Rome.

1. Introduction
It is no exaggeration to say that the Asia-Pacificregion is a treasure house ofanimal genetic resources. The percentage of worlds animal species and breeds in thisregion is 32 % for cattle, 96% for buffaloes, 56% for pigs, 40% for sheep, 57% for goats,
43% for chicken, 83% for ducks(Sasaki,1995).
Furthermore, according to the Global
Data Bank for Animal Genetic Resources of the Food and Agriculture Organization of
the United Nations (FAO), the total number of farm animal breeds which were registeredis 2,719 in the whole world and 746 breeds (about 27%) are located in the Asia-Pacificregion (FAO, 1993b).
The global environment has altered as a result of human activities,
developmentof social and economic system and rural development. As a result, the number of theanimal species on earth, including wild life and domestic animals, is decreasing rapidly.
This trend is marked in the developing countries including those of the Asian region. The
population of many pure indigenous breeds is likely to diminish as a result of crossbreed-216

ing, replacement or neglect. It is supposed that many indigenous species are at risk ofextinction.
FAO started a new regional project entitled "Conservation and Use of Animal
Genetic Resources in Asia and the Pacific". This project is supported by the Japanese
government. The author was sent to the FAO by the Japanese government andparticipatedin the activities of this project for 18 months. Based on the experience ofthat period, the author introduces the activitiesof the project, with special focus on theinformation system for the animal genetic resources in the Asian region.
2. The recent international
movement on animal genetic resources
Before discussing
the main issues, it is useful to understandthe recent
international
movementon animal genetic resources.
Since its establishment,
FAO has been involved in the conservation of the animal
genetic resources. During the 1 980's, FAO implemented many activities in cooperation
with the UNEP (United Nations Environment Programme). In 1989, the Committee ofAgricultureof FAO, which was held in Rome, Italy, discussed
Animal Genetic
Resources. The Committee reviewed the past, present situation and the future prospects
for animal genetic resources activities.
The Committee recommended orientation and the
problems for FAO and for the national governments Afterwards, the FAO held twoExpert Consultations on Animal Genetic Resources: (a) A global program for sustainable
development and (b) Management of Global Animal Genetic Resources in 1989 and1992, respectively.
The programmes and activities
were designed and implemented
based on the recommendations of these consultations.
During the same year, the decisionof the United Nations Congress on the Environment and Development (the "Earth
Summit"), which was held on Rio de Janeiro in Brazil in June 1992, and the Convention
on Biological Diversity, which was ratified in December 1 993, gave recognition to thenecessity of global scale activities
on conserving biodiversity.
At the same time, a newperspective was added that of sustainable agriculture.
Based on these developments, theFAO developed plans for the conservation of domestic animal diversity and is in the firststage of these activities.
Global Management of Animal Genetic Resources was
discussed
at the Committee of Agriculture of the FAO held in March 1995 in Rome,
Italy. Based on the decision of this Committee, FAO recently developed a Special Action
Programme for the conservation of animal genetic resources. The Programme intends to217

Table 1. Number of Animal Breeds collected
B u ffa loC attle Y a k H o rse A s s
C h in a
L ao sM a lay sia
M yanm ar
N e p alT h a ila n d
V ietn a m

5
l
2
2

9
l
4
5

5

10

2

‑

‑

‑

1
l

3
l

‑
‑

1
1

‑
‑

T o ta l

17

30

6

13

by the ProjectG o at S h ee p P ig C h ic k enD u c k G e e seT u rk eyC a m e Rl a b b itD e e r T o ta l
4

24
1
1
3

1

1
2

2 1
1
1
1

3

1
1

1
‑

3
5

2
7

16

30

40

ll

83

1

4
3

1
1

1

1

13
28
17
22

1

1

18 1

establish
as quickly as possible a sound primary infrastructure
for conserving
genetic resources which will involve many countries and organizations.

animal3. Special

Action Programme for the Animal Genetic Resources
FAO designated a number of imperative actions for the successful managementof animal genetic resources (Hammond, 1994). These actions include:
1 ) identify and understand unique resources2) develop and properly use these resources3) monitor the status of the resources at risk
4) preserve those unique resources which are not currently in demand5) train and involve national scientists6) communicate to world opinion the importance of the sustainable
management of animal genetic resources.
Furthermore, the FAO developed a Global Strategy for the Management of
Animal Genetic Resources. This Strategy includes the following components :* An intergovernmental mechanism* A geographically
distributed
global structure* Activities,grouped into seven work elements* Expert panels ensuring sound advice for the Strategy's implementation
The specificfeatures as follows :1 ) Execution

activities

to execute this programme are composed of some keyof regional

project

identification

missions

to promote effectively

218

2)

3)

4)
5)
6)
7)

8)
9)

many conservation activities
Establishmentof geographically
distributed
structure
To establish this structure, it is necessary to build the components as follows:(1) umbrella projects in each region of the world in order to involve andassist governments in designing and implementing national action plans(2) a coordinating
institution
within each country with an active country
coordinator(3) a global focus in FAO Headquarters in RomeFormation of a Domestic Animal Diversity Information System (DAD-IS)
To form DAD-IS, it is necessary:- to develop rapidly the Global Data Bank on Animal Genetic
Resources- to expand rapidly the World Watch List for Domestic Animal
Diversity
Fostering an in-situ conservation strategy
Introductionof an ex-situ conservation strategy- Establishmentof national and international
gene banks
Monitoring of populations of breeds at risk
Implementationof a specific global research activity- To establish the amount of diversity in each domestic species, andthe contributionof breeds to diversity- To study a genetic distance- To make genetic maps
Implementation of the activity for the communication of the information andthe training of the experts
Development of the guidelines
to establish
a plan for the international
activities
and the national activities

4. The project of "Conservationand Use of Animal Genetic Resources in Asiaand the Pacific"
This project is one of FAO's trust fund projects supported by the Japanese
government. The Japanese government provides US$1.82 million to FAO and FAO
carries out the project activities.
This project is coded MGCP/RAS/144/JPNM.
The

219

duration of the project is for 4 years from December 1993 to December 1997 and covers
1 2 Asian countries. The participating
countries are: Bhutan, China, India, Indonesia,
Laos, Malaysia, Myanmar, Nepal, Pakistan, Philippines,
Thailand and Vietnam. These
countries covers over 85% of the animal genetic resources in the region and therefore
over 25% of the global resources.
The main activities are:1) To identify and monitor the animal genetic resources of the 12 countries
To make a regional inventory of all known breeds, breeds varieties andpopulations for all livestock species (buffalo, cattle, horses, pigs, sheep,goats, poultry, ducks, deer, yak) with information on population sizes,
as well as on evolution of populations
and also with some performancedata. For two countries , it is a first time to do an actual breed survey.
2) To preserve and simultaneously enhance the productivityof those indigenous
breeds at risk of disappearing.
3) To train national experts on techniques for descriptionof breeds, data
handling and in-situ and/or ex-situ preservation.
4) To analyze and publish
all information
related with Animal Genetic
Resources collected or created during the project.
5) To establish
the Asian Network on Conservation of Animal Genetic
Resources.
The initial activities
are concerned with collecting the latest population data oneach breed in each country. The project office is located in the building of FAO's
Regional office for Asia and the Pacific (RAPA)in Bangkok, Thailand. The staff of the
project are a Chief Technical Adviser sent by FAO Headquarters, a Senior Animal
Resources Officer sent by Japanese government (the author) and a secretary employedin Bangkok.
National Coordinators were nominated and decided in each of the 12 countries.
They are responsible for the project activities
in each country and the institution
at whicheach works is the Coordinating Institution
for that country and will be the center for thewithin country network.
5. Establishmentof the Network on Animal Genetic Resources in Asian region
As mentioned above, the establishment of the Network is one of main objectives

Figureof the project

1. The model structure

MGCP/RAS/144/JPN".

of the Asian network

This is the first stage in the global

programme of

FAO.

To understand the concept of the Network, it is useful to understand the Global
Data Bank for Animal Genetic Resources at FAO Headquarters in Rome and World
Watch List for Domestic Animal Diversity.
1) The Global Data Bank for Animal Genetic Resources
With the help of many organizations and individuals
throughout the world,
FAO initiated the global breed survey of main domestic animals in 1 99 1 and establisheda Global Data Bank for Animal Genetic Resources at FAO Headquarters in Rome in1992. This Global Data Bank will have a core role in the Global Network. The Data
Bank now includes initial data on about 2,800 breeds (and varieties of breeds) of 7
species (ass, buffalos, cattle, goats, horses, pigs and sheep) and population size data for
about half of these breeds. It is planned to include camels and poultry and the survey ofthese two species is progressing now.
The list of breeds is still in the development phase and it is certain to change as
information accumulates. Population data are currently entered into the Global DataBank.

For each breed, the following information are entered ~ its origin, population
size, physical and morphological features, uses and special attributes, production records
and managementconditions - in addition to information which is less commonly foundsuch as genetic distances and genetic conservation programmes for the breed. The world
dictionary of livestock breeds by Mason (Mason, 1 988) was used as the primary sourceof information on breeds and breed varieties for the data bank. This book provides breed
names and synonyms, the geographicallocation of the population as well as giving abasic description of the origin, physical appearance and main uses of each breed.
2) The World Watch List for
Domestic Animal Diversity
Together with UNEP, FAO
published
the first edition of theWorld Watch List for Domestic Animal Diversity in 1993 (FAO, 1993b).
This list will act as another component of the global early warning system for domestic animals and wascalled for under the Convention on
Biological
Diversity.
The early results suggest that at least 30% andpossibly
as high as 40% of all animal genetic resources are currently athigh risk of extinction.
The simple
criteria used to determine the risk ofextinction
was, less than 1,000
breeding females and less than 20
breeding males remaining. The majority of the breeds at risk are indeveloping
regions.
Adequate records do not exist to enable us toobtain a reliable estimate of either
loss
rates
of these
breeding
resources or of domestic animal diversity itself.
The second edition
will be published
soon.

222

Regional

Center* National

Figure3. The basic model structure

Centerfor DAS-IS.

3) The Network on Animal Genetic Resources in the Asian region
The regional center for the Network is located in the office of the project"GCP/RAS/144/JPN"
which is in the RAPA. The Asian databank is also located in the
project office. This regional databank has a core role in the Network. The institutes,
inwhich the National Coordinator works in each of 12 countries, will be a national centerand the national databank will be developed there. The national databank is also essential
for the national Network and it will also have an important role in the Asian Network.
In these institutes,
E-mail systems are being set up and these institutes
will be linked to
the regional center in Bangkok. The model structure of the Asian Network is shown(Figure 1) and the model structure of each country is shown (Figure 2). The regional
center is linked to the global center in Rome. These structures are a component ofDomestic Animal Diversity
Information System (DAD-IS).
Namely, DAD-IS iscomposed of a Global center, Regional centers and National centers. DAD-IS will be
operated through Internet based on a Personal Computer. DAD-IS will serve as an
information axis for the FAO's Special Action Programme and as a global early warning
system. DAD-IS will also provide easily available databank, and have research design
and analysis, training and communication capabilities.
The basic model structure for
DAD-IS is shown (Figure 3).
Each country should organize its own databank according to its circumstances.

Figure4. White Lamphun Cattle

Figure6. Tre Chickenin Thailand.

in Vietnam.

Figure5. Ac Chicken

Figurein Vietnam.

7. A yak in Nepal.

224

Once a national databank is established,in principle, each country should take care of theconservation of its own animal genetic resources. The countries participatingin theproject will be responsiblefor decisions concerning the breeds/populations
to beconserved. They will be influenced in these decisions by some guidelines commonto all
countries involved in this activity.
Conservation of animal genetic resources must be anational initiative.
Given the substantial differences in needs and appropriate approaches
in individual
countries, the organization of national conservation will vary widely. It isalso important that private and public interests within in each country are incorporated
into the strategy for conserving animal genetic resources to ensure an effectively,
coordinated national program. The support of strong national programs is essential to
the success of regional and international
conservation efforts. Ultimately,
livestockproduction and improvement programs will benefit from international
support andassistance from enhanced international movement of animal genetic resources.
It is necessary to create an appropriate infrastructuresin each country. New
institutional
facilities
to support field activities will be needed. They need not be large,
but they must be suitable for liaison with national governments, international
bodies bothgovernmental and NGO's (Non Governmental Organizations^),
and with bilateralorganizations
working on animal genetic resources. They should include Cryogenic
Animal Gene Bank(s).
6. Some information on the animal genetic resources in the Asian region
The animal breeds collected by the project by the end of June 1995 is shown(Table 1). The number of the animal breeds collected is 1 8 1. This number includes only
7 countries, and for 3 countries this does not include all of the breeds because the breed
survey is not yet complete. These data were sent to FAO Headquarters and were entered
to the Global Data Bank for Animal Genetic Resources.
The project collected information on the breeds at a serious risk in each country.
And the project decided to provide budgetary support to some of the breeds which needs
urgent action for conservation. Some of the breeds are as follows:
1) South China Pig in Malaysia
2) Shwe Ni Gyi Cattle in Myanmar
3) White Lamphun Cattle in Thailand (Figure 4)4) Ac Chicken in Vietnam (Figure 5)

225

5)

Tre Chicken in Vietnam (Figure 6)
The yak in Nepal, is not endangered but it is one of very particularshown (Figure 7).

breed in Asia, is

7. Conclusion
This project is a first umbrella project for animal genetic resources in the world
and it is conducted as a Special Action Programme of FAO. This project is also being
used as a pilot scheme for the global programme. It is significant that this project wasstarted in Asia. The author wishes the project success in attaining its objectives.
References
FAO. 1990.

Animal genetic

resources-A global

Animal Production
FAO. 1992.

and Health

Healthanimal genetic

globalDivision,

Publication,

Conservation

R. 1994.

Introducing

1988.

World

Ruane,

J.

region.

1993.

Applieddictionary

to Livestock

to Livestock
Meeting,
Yamada, Y. 1995b.
Coordinators

Global

Production,

breeds,towards the year 2000-Overall
Proceedingsof the First NationalandEds. R. Loftus and B. Sherf,

overview. Proceedingsof the 5th

2 1 : 423-430.
animal diversity.

Animal Genetic

Types

and varieties.

Third

edition,CAB

the world's

domestic

review of livestock
Coordinators
animaldevelopment

Meeting,resources.

in the Asia-

FAO Bangkok, p4-6.

AnimalGenetic

Resources

FAO, 1 1 : 13-21.
Current development-the Asia and Pacific Region.

Yamada, Y. 1995a.

1st Edition,

animal diversity,of livestock

Steane, D., Chen Y. and Yamada Y. 1994.
diversity

Production

UK.

Documenting

Information,

AnimalFAO, 13: 3-1 1.

Wallingford,

M. 1995. Challenge
Pacific

among the breeds ofof a Working Group, June 1993.

the first world watch list for domestic

Resources Information,
International,

relationships

Rome, Nov. 1 993.

of domestic

World Congress on Genetics

Sasaki,Reports

the genetic

World Watch List for Domestic Animal Diversity,

Hammond,K. 1994.

Mason, I.L.

andFAO Rome.

Joint FAO/UNEP

Loftus,resources. Ed. J. Hodges, Animal Production

programme to establish

each domestic animal species.
FAO. 1993b.

Ed. G. Wiener,

Paper, 1 04, FAO Rome.

An integrated

Healthdevelopment.

Paper, 80, FAO Rome.

The management of global

FAO. 1993a.

programme for sustainable

Production,

Proceedingsof domestic

animalon Genetics

Applied

21 : 23 1-238.

Questionnaire

for breed

survey. Proceedings

FAO Bangkok, p8-1 3.
Network for conservation of animal genetic
Meeting,in the conservationof 5th World Congress

FAO Bangkok, p i 4-36.

resources.

of the First National
Proceedings

Coordinatorsof the First National

227

Questionsand answers in Session 5

Comment from Dr. Mikami
Q. In Japan where are per primers available?
(Astuti)
C For cattle, the Shirakawa Institute of the Livestock Technologal Association provides
more than 200 per primer sets for MS markers. In pigs, the National Institute ofAnimal Industry and STAFF Institute also provide about 200 primers sets. We are
studying linkage analysis between these markers and economic traits using large
resources (reference)family produced by crossing between geneticallydivergent
breeds, for example between improved breeds and native breeds. (Mikami)
Question to Dr. Yamada
Q. When you mention collected breeds from Asian countries do you meandata or live
animals? Why are Indonesia and the Philippines
not on the list? (Matias)
A. Collected data on breeds. By the end of June we had not received the questionnaire
sheet from those two countries. (Yamada)Nepalese

Animal Genetic Resources(Paperat the MAFF Seminar)

presented

NANDAPRASAD SHRES THA
Animal Breeding

Division,

Nepal Agricultural

Research Council

P.O.Box 1950, Kathmandu, Nepal
Summary
Nepal is a small mountainous Himalayancountry with an area of 147,181

people.

The livestock

population

millionsheep, 5.452

milliongoats, 0.605

million

Nepal has diverse

ecosystems

and diverse

Nepal has different

in Nepal comprises

types of wild animal genetic

deer, Jungle fowl, tiger, elephant,in natural

habitatrhinoceros,of 6.234 million
pigs,

cattle,3.073

km2 and 18.5 million
millionbuffaloes,

and 0.392

millionmillionpoultry,

genetic

resourcesof plant, animals and other species.

resources such as Gauri Gai, Arnii buffalo,deer, bear, wolves. Nepal has varietiesthat are commonto subtropical

0.91 113.601to temperate

ducks.

wild pig, muskof plantsthat are foundand alpine type of climate

and Himalayan

ranges.
Nepal is rich in domesticatedresources
supplycontributeabout one/third

animal geneticof the National

diversity

also.

Agricultural

The domesticatedDomestic

animal genetic

Gross Product.

milk, meat, draft power, manure, hides and skin and wool as the product

The livestock

sources of livelihoodofmanypoor Nepalese farmers. The contributionof livestock in the high hill and mountain is higher than in
Terai Region where crops dominate the socioeconomic condition of the farmers compared to the hills andmountains where the land available
Distribution

for crops is scarce.

and availabilityof different

present

in term of evaluation

is described.

Theircharacteristicsof the available

statusbreeds of buffaloes,

breeds is discussed.

cattle,of the production

The Lulu cattle,

AchhamiPate breed of pig and Kage sheep may become extinct soon if suitableconservation

as the number is decreasing

rapidly

Yak/Nak,measures are not taken.

They need

as a result of crossbreeding

Natural selection has been operating for many generations
in considerable

Physicalpopulation

hardy condition

barriers

diversity

not well identified,and replacementby exotic

animals is low compared to someon the livestock

to movement of animals isolatedin Nepal.

characterized

Many breeds and strains

and evaluatedfor exploitation

populationin differents

for many generations

has resulted

exist withineach species

particularly

for traits

and they arelike resistancetoand diseases.

Buffalo is the main livestock

species contributing

type called the Lime and the riverine type called the Parkote
In addition,and other

breeds.

parts of the country.
stilperformances

Cattle, Lime Buffalo,

breeds. The economic performance of these native breeds of domesticatedimported

sheep, goat, pig, ducks, poultry

Terai and other uncharacterised

breed which is spreading

about 52 % of the livestock

are the two types of local buffalo found in Nepal.

types are also found in the country.

all over the country for both crossbreeding

milk and meat production.

The Swamp type of buffalo

GDP. The Swamp

Murrah are an imported

purposes and to keep as a pure breed forneeds conservation

as this type may be lost if they

230
are not protected.

The wild buffalo

Lulu, Siri, Achhami,

Arna is being conservedand Hill cattle

and Terai Cattle

The Lulu, Achammi and Siri cattle need immediate attention
rapidly.

in the Koshi

Jhyopkyos

Park.

are the five types of cattle

found in Nepal.

to conserve them as their numbers are declining

The Gauri Gai is been conserved in Chitawan National
The Yak/Nak, Chauries,

Tappu NationalPark.

are reared in the high mountains

and Himalayas.

They aremulti purpose animals for milk, meat, draft power for plowing, transportationof goods in high mountain on
small tracks. Their numbers are decreasing rapidly and they are in need of characterization,
evaluation, andgenetic

improvementby adopting

a farmers group breeding

approach.

The goat meat is accepted by all the Nepalese people. Goat meat is in always in high demand. The
goat population is increasing.
Chyangra in the high Himalayas,
Sinhal in the high mountains, Khari at mid
hills,and Terai goats are the four types of goat based on the ecologicalbeen subjected

to cross breeding

Bhyanglung,goats from India particularly

The goat has

Jamunapari

and Barberi.

Baruwal, Kage, and Lampuchhre are four breeds of sheep occur in Nepal.

breed has been decreasing
Sakiniwith the imported

system of the country.

at fast rate and soon it may extinct

and Naked Neck are the poultry

if proper

The Kagemeasures are not taken.

type breed found in Nepal.

scavengers. They have been cross breed with New Hampshire and Australorp.

They are kept as backyard
Some Nepalese Red Jungle

Fowl is found in forest areas.
Hurrah and Chwanche are the two types of pig found in the Terai and Hills

of Nepal.

are also found in the Terai and High Mountain areas. The wild pig needs conservation.
Hurrah breed could be exploitedfor their

During the last three decades
and propagationof imported

Wild pigs

The Chwanche andsmall size and early maturity.
the Government of Nepal has been giving

animal breeds of different

domesticatedspecies.

emphasisto production

Native breeds were not given

due consideration compared to economically competitive imported breeds. Some awareness has been createdby the organization of a Workshop on animal genetic resources conservation in Nepal' by joint collaboration
between Nepal Agricultural
Breeding Division,
ll-13 April 1994.

Research

Department

The Nepal Agricultural
Research including
biodiversity.

bio-diversity.

Council,of Agricultural
Research

Development
Councilthe research capability

facilities

in relation

is not done in Nepal either for plants or animals.

AnimalScienceResearch Institute,and United Nation Department

is the coordinating

The crop sector of biodiversity

The lack of manpower and laboratory

a focus on increasing
technical

National

national

insitute

AnimalProgramme,for Agriculturalis at a more advanced stage than the animal
are the main constraints

and there needs to be

to animal biodiversity

research.

Genome analysis

Foreign Research Institutions

supportboth financial

andwill be needed to carry out such research work.

Introduction
Nepal is a small Himalayan country with a population of 1 8.5 million and areaof 147. 181 km2. Nepal is blessed with the worlds highest mountains, deep gorges, fast
flowing rivers and gorgeous valleys.

Despitethe small size of the country, Nepal has a231diversity of ecosystems, animals and plants. The tremendous differences in ecology andenvironment within a short distance are mainly due to big changes in mountaintopography,
isolated valleys and niches and varying hill slopes and aspect. Such
topography has generated tremendous bio-diversity
in plant and animal genetic resources.
Basically the whole country can be divided into five geographicalregions. Trans-Himal/High Himal, High mountain, middle mountain, siwalik/ lower hills and the Terai (Figurei).

Animal Genetic Resources Across Different Ecological
RegionTrans-Himal/High Himal Range (Figure 1)
This regions is characterized
by the area situated at higher than 2500m andadjacent to the Tibetan border behind the Himalayan mountain ranges. This area ischaracterized by high peaks, steep slope, narrow valleys and gorges. The climate is arid,
dry cold and windy. The rainfall ranges from 200-800 mm/annumwith intense solar
radiation, high winds and high evaporation rate.
The people living in the high Himalayan areas are Sherpas in Solukhumbu,
Thakalis in the Dhaulagiri,
Bhotias in Humla, Dolpa and other areas bordering Tibet.
They are involved in the barter system of agricultural
products with Tibetan neighbours.
The barter system has been disturbed recently due to easy access with India. For the
Serpas their lifestyle has changed, many are tourist guides or camp helpers.
The major vegetation of this region consist of dwarf shrubs with some perennial
pastures and weeds.
The major domesticated animals raised in these areas are Chyangra goat, Lulu
cattle, Yak Chauries, Kirko, and Bhyanglung sheep, Tibetan horse and donkey. About
3% of the total Nepalese population live in this region. Potato and buckwheat are themajor food crops.
The animals are kept in a nomadic migratory system herds are taken to higher
altitudesin summer and lower altitudes in winter. Previously farmers were taking
animals across the Tibetan border.
High Mountain Range (Figure 1)
This range lies at an attitude of 2200-4000m
lying south of the Himalayan
ranges. The climate and vegetation varies from temperate to alpine. High peaks are232covered with snow in the winter and when the snow melts these change to alpine pasture.
The major feed resources for animals are from alpine meadows and shrub forest cover,and somefromthe crop land. The major animals raised in this region are Yak, Chauries,
Kirko Cattle, local hill cattle Baruwal/Dhorel
Sheep, Sinhal Goat and hill buffalo. About
8% of the Nepalese population live in this region.
Animals are reared in a nomadic migratory management system taking theanimals to alpine grazing areas in summer and to lower altitudes in winter. Animals areused to transport produce from Tibet and from the Terai during summerand winterrespectively.
The sheep and goats in the mid western region are also used for carrying
food grain from the Terai and salt from Tibet.
The people residing in this area are Tamang, Limbu, Gurung, Thakali, Bhotias,
Magar. They live in a poor environment by using the natural resources of the region.
Their main livelihood
is from livestock supported by some crops such as potatoes,
buckwheat, wheat, millet that are grown in limited areas.
Mid Mountain Ranges(FigurQ I)
This Zone represent the areas ranging from 800- 2400m lying north of Siwalik
ranges. The climate and vegetation is mainly sub temperate and temperate with aconsiderable
variation in micro environment.
This region is the mostly densely populated with about 46% of the people livingin this region. Bhrahman, Chhetri, Gurung, Magar, Newar are the main dwellers of thisregion.
Commonpractices of rearing animals in the region include a semi nomadic
migratory system, stall feeding and taking the animals out to graze in the day time andkeeping in sheds at night. Some intensive animal production systems are commonin
areas surrounding city areas.
The major feed resources come from grazing on high meadows, shrubs,
communal grazing lands, waste lands, and forests and a limited amount of crop by
products.
Different type of fodder trees are commonin this areas for feeding animals
during the lean period.
These major animals raised in this region are Hill cattle, Achhame cattle, Siri
Cattle, Baruwal and Kage sheep Sinhal Goat, Lime and Parkote buffalo, local black pig(Chwanche) Sungur, Pundi and Sakini and Naked Neck chicken.

233

FigureSiwalik

1. Map of Nepal. 1) Trans-Himal/High
Himal Range (2,500 - 8,848 m), 2) High Mountain Range(2,000 - 2,500 m), 3) Md Mountain Range (700 - 2,000 m), 4) Siwalik hill (300 - 700 m) and
5) Terai Region (below 300 m). Bar = 200 km.

/ii//s(Figure
1)
This region includes mainly low land and upland forest and agriculturalareas
ranging form 200 m to 1500 meter elevation.
The climate is sub-tropicaltosub-temperate. The major feed supply comes fromforest grazing, crop lands and waste
lands. Most of the animals are raised under sedentary management system based on thegrazing in the surrounding forest, fodder trees, agriculturallands/fallow,
waste land,terraces and rivers.
Animals are kept in sheds during night and are taken to grazing areas during theday. Stall feeding the milking animals is also common.Paddy straw is the most common
feed for stall feeding, as well as, additional
feeding during lean period of available feeds
form grazing land.
The Tharu and the migrants from India and hills and mountain areas are thepeople living in the regions.
The genetic resources available in this region are Kage sheep, Khari goat,
Chwanche and Pundi pig, lime buffalo, Pahadi hill cattle. Some exotic breeds and theircross breed with buffalo, sheep, cattle and pig are also raised in this region.

234

Terai Region (Figure 1)
This region is the plain situated below 300m and is the main crop growing areaof the country.
The Tharu and migrants from India and different regions of Nepal live in thisregion. The region is the main grain production center for the whole country.
The major feed supply comes from cropland residues and grazing fallow land,waste land and roadside verges. Growing of fodder crops is increasing in the region.
The animals are kept in sheds during night and are taken out for grazing during
the day. Feeding animals with paddy straws is quite common.Intensive stall feeding of
milking animals is commonaround markets and cities.
The major animal resources in the region are Terai Cattle, Terai Buffalo Terai
Goat, Hurrah pig and Lampuchhre sheep. The cross breeding of Terai cattle with Jersey,
Holsteins Friesians, and Terai buffalo with Murrahs is increasing.

Brief Descriptionof Different Breeds of Different Animal Types
Buffalo Genetic Resources
Wild buffalo or Arna (Bubalus arnii) are seriously endangered and have areconserved in Kosi Hill Tappu riverine terraces at 600-2000ml.
Their morphologicalcharacteristics
are rounded for head with large horn and dark black body color. They
measure 3.2 m long frommouth to root of the tail and 1.8 m high at the shoulder. Little
is known about production and reproduction performances of these buffalo. Genome
analysis, electrophoresisand other studies will be areas to conduct future research.
Domesticated buffalo provides about 5 3% of the livestock contribution to thenational GDP and about 71% of the total milk ( 87,700 tons) and 64.4% of total meat.
About 12% of the farmers use buffalo as draft animals for ploughing and for transporting
agricultural
products. Two type of buffalo are found on Nepal, the riverine and theswamptype. The swamp type buffalo differs from riverine geneticallyin color and body
appearance.
Lime:
Nepalese swamp-type buffalo {Bubalus bubalis)are believe to have been
introduced to Nepal long time ago from China. They are found in the mid hill and highhill region of Nepal. The animal is characterized by chevrons of gray or white hair below
the Jaws and under the chest. Its Sickle shaped horn curved towards the neck. The

235

populationof lime is decreasing very fast through indiscriminate
crossbreeding
withriverain type. Pure males of this type are difficult to find these days. This breed has poor
milk production and has been replaced by riverain buffalo.
Parkote: The native riverain type buffalo are mainly in the lower hills and Terai. They
have not been characterized.
They are used to cross with Murrah breeds for milk
production. Parkote are medium size breed slightly bigger than lime and smaller than terai
buffalo.
Non-descript buffalo: There are many non descript buffalo in Nepal some of these havebeen introduuced from India. They have not been studied well. They have not been
evaluated for the production, reproduction and other parameters.
Imported buffalo: The main imported breeds are Murrah. Some NiliRavi types can beseen in some places.
Cattle Genetic Resources
The wild cattle, Gauri Gai (Bos gaums) are found in Chitawan National Parkand they have not been studied for their production, reproduction and other parameters.
About 7.3 million domestic cattle are reared by 2 million Nepalese farmers. The
cattle population comprises 3.2 million adult male mainly for ploughing and 2.3 million
adult females which provide 30% of the total milk supply (876,000tons).
Terai white zebu cattle:
Most Terai cattle have white hair and black skin.
Crossbreeding with Haryana Cattle has been practiced for draft purpose. The height atwither is about 141 cm for bulls and 131 cm for cows. Body weight of bull is about360-500kg and of cows about 3 5 0kg. For production and reproduction parameters thisbreed needs to be further studied.
Black hill Cattle: Diversified types of black hill cattle exists in Nepal. They have been
kept isolated for long time due to topographicalbarriers.
Siri Cattle:
According to Epstein ( 1977) Siri Cattle were introduced to Nepal through
Darjellingfrom Bhutan. The few Pure Siri Cattle that exist in Illam may become extinct
soon. However some crossbreed Cattle are found in Illam district.
The Siri cattle areblack and white in color with well developed udder. They are larger than the black hill
cattle. The neck is fairly short and dewlap is moderate in size.
Lulu Cattle: The humpless cattle are reared in Mustang district. Similar type of cattle
are also found in Solukhumbu area brought from Tibet. Their number is decreasing236

sharply.
Achhami Cattle:
This cattle being small in size are known as " Sano Gai " meaning
dwarf cattle. They are few in numbers and are not well characterized.
The pure form of
Achhami Cattle is difficult to find and they are decreasing in number because ofcrossbreedingwith other breeds.
They need to be characterized
and identified forphysical, production and reproduction performances.
Imported breeds of Cattle:
The Jersey, Friesian, Brown Swiss and Ayreshire are themilk breeds that were introduced to Nepal. They have been used for their semen. The
Haryana Breeds have been used for cross breeding with the terai cattle for draft purpose.
Yak Genetic Resources
Yak {Bos Grunniens) are kept in the high Himalayas where they are well adapted
to severe cold and can withstand lack offood for a number of days. Yaks are found in
Solukhumbu, Jomsom, Dolpa, and Mugu but have not been well studied because of
remoteness and difficulty of access to the rugged terrain. They descend only to 4000m
above sea level in winter and graze at 6000-7000m during the summermonths. They are
reared as a transhimalayan
animals. These multipurpose animals have been used formeat, milk, ploughing, and for transporting goods over small tracks in the high mountain.
The number of Yak/Nak is decreasing very fast. Both Yak and Nak has been used forcrossbreeding
purposes to produce Urang Chauris (Jhoopkyos)
and Dimzo Chauris(Jhypkyos).
The gene flow fromthese crosses stopped because no further generation are
produced after the first and second crosses as they are sterile and are selective to breed
the female with either Lang or hill cattle.
Milk protein polymorphism in Yak/Nak cattle and their hybrids were examinedelectrophoreticallyby Yoshi Kayoto and others from the University of Tokyo in 1 988.
The result suggest that gene flow from high mountain to low areas of Nepal is low. The
genetic constitutionof Yak was estimated to be much differentiatedfrom breeds ofEuropean and Indian cattle.
Goat Genetic Resources
The goat represent the largest population of small ruminants in the country.
They are distributed
all over the country. Different breeds exist in different ecologicalareas. The management system also varies with the elevation. The various types of goat

237

raised in the country are as follows:
Chyangra: This type of goat is reared in the Himalayan ranges of the country. It is well
adapted to cool, dry, semidesert snowy type of climate. The Chyangra found in Nepal aresmaller than those of Tibet. This type of goat is known as Kasmiri in some areas andChyapu in other areas. The male weighs about 45-60kg and the female weighs about
35-50 kg. Each she goat produces one kid per year and about 125-200gm at Pasmina.
The have not undergone any artificial selection and have been reared in the yak rearing
ranges. They have not been studied well because of remoteness and difficulty of getting
to areas where they are reared. They are mainly reared by the Humli, Dolpali and Sherpa
people. They are decreasing in number because of restrictions in the grazing areas ofTibet.
Sinhal:This high hill goat is kept along with flocks of Baruwal sheep in the ranges of500- 1500m above sea level. Male buck weighs about 40-50kg and the female weighs
30-40 kg. Both male and female have horns and medium sized ears. The goats are
generally of mixed color black and white or pure white or pure black. This breed is being
indiscriminatly
crossbreed with Chyangra or Khari breeds. Their number are decreasing
rapidly.
Khari breed: This breed is reared in the lower mountain areas. These are smaller thanother breeds of goat. Males weigh 30-40kg and female weigh 25-30kg. Both sexes havehorns and the coat is multicolored.
They are a prolific breed and twinning is a common
trait. The frequency of breeding is also high and three pregancies in two years is possible.
Sometimes 4 kids per kidding can be observed in farmers flocks.
Terai goat: They are long earred and are of mixed type. They resemble Jamunapari.
They are bigger in body size with body weight of male 40-50kg and of female between
30-40 kg. Other imported breeds of goat are Jamunapari and Barberi from India. A few
crossbreeds with Sanen are also available.
They are a good milk yielder. The crossbred
with Sanen have a higher milk yield than other breeds in Nepal.
Sheep Genetic Resources
Four different type of sheep are found in Nepal. They have undergone natural
selection only. Their genetic variation seems to be very high.
Bhyanglung:This breed originated in Tibet. They are reared in the border areas with
Tibet. They are kept at altitudes in excess of 3000m. They are adapted to the high cold

238

desert type of ecosystem where they live. They cannot survive on lower hills. Adult
males weigh between 30-45 kg and females weigh 25-35kg. Their wool is very good for
Nepalese carpet making blended with New Zealand carpet wool. Their wool is finer thanthat of other breeds. They have not been studied for their production and reproduction
characteristics.
Baruwal Sheep: This breed is a hardy mountain breed. Males weigh between 35-45kg
and female weigh between 30-35 kg. Their legs are long and strong and are suitable fornomadic sheep rearing. They can run fast in the poor hill and mountain environment.
They are also used for transportationof goods from the Terai to mountains and from
Tibet to lower hills. The heads are large with small ears. The wool is generally white but
some have black and brown patches on the coat. Rams have curled horn and ewes arehornless or with smaller horns. They are kept mainly for meat and wool in the westernregion and also for draft poposes in the mid-western region.
Kage Sheep: Kage sheep are a valley breed found in the mid mountain and hilly regions
ranging in elevation from 600m to 1500m. This is the smallest breed of subtropical typeand are generally polled females. Adult males weigh between 20-25kg and females 15 -20
kg. The wool yield is about 300-500g per year per sheep. They are non-seasonal
breeders.
Twinning is commonin the Kathmandu valley while twinning is rare in the
Pokara valley. It is now difficult to find pure male kage sheep because of indiscriminate
cros sbreeding.
Lampuchhre Sheep: This breed resembles kage type but has mixed color on the neckand head and is much bigger than Kage with long tail and is raised in the western Terai.
The body is generally covered with coarse white wool.
Imported sheep breed: Polwarth, Merino, Romney Marsh, Border Leicester, andRambuillet have been introduced for crossbreeding.
Pig Genetic Resources
Most of the 496,000 pigs in Nepal are the native type. They are reared by theTharu, Magar, Limbu, Damain, Sharkee people of the country. Many breeds have notbeen characterized
and identified
for their phenotypic,
production and reproduction
performances. Chwanche pigs are found in the western region in the Magar community,
Sungur is found in in the mid western and western region, Pundi is found in the easternregion, Hurrah or Pate is found in the Terai region. These breeds are raised on a small

239

scale in a backyard scavenging system with a small amount of kitchen waste, grain
by-product, and other type of waste as feed. They are raised with almost no veterinary
care and are supposed to be disease resistant to many diseases and parasites.
They
become sexually mature when they are about six months of age.
Somewild pigs are also available in the forest ranging from Terai to Jumla areas
to an elevation of about 3000m. They have not been studied thoroughly.
Some of the imported breeds of pigs in Nepal are Yorkshire, Landrace,
Saddleback
Hampshire, Tameworth and Duroc. Their production
performancescharacteristics
in Nepal are being evaluated.
Horses, Mules andAsses
There are about 19,000 horses, mules and asses in Nepal. Mules are morecommonin Western Nepal where they are mostly used for transportationof goods in thehigh mountain areas. The different type of horses found in different parts of Nepal are
namedafter the respective districts.
Namely Mustangi in Mustang, Jumli and Humli in
Jumla and Humla districts respectively.
They are believed to have originated in Tibet.
However these horses are smaller than the Tibetan type. The number of asses are notknown.
Blood protein polymorphisms of Tibetan asses in Nepal were examined andsubstantial genetical differences between asses and horses were reported by Kawamotoetal (1988).
Poultry Genetic Resources
The Sakini type and Naked neck type of poultry are available in different partsof the country. They are small, their eggs are small, and characterized
by broodiness offemales. They are given no health care and are raised in a scavenging system withhousehold waste and free grazing around the houses. They are resistant to many parasites
and manydiseases and can withstand poor environmental conditions.
Their meat is said
to be tastier than the broiler commercial type. Naked neck are said to be meatier than thefull feather type. They are said to be more suitable to warmer climates.
Duck Genetic Resources
Muscovy ducks and species

are found in Nepal.

240

Genomic Analysis
Electrophoretic
study on blood protein and milk protein of yak, cattle and theirhybrids and blood protein of asses, mules and horses of Nepal and of native goats werestudied by Takao Nishida and other investigators in 1 988. The same group made a studyof different breeds of sheep for the genetic distances and similarity
by taking blood
sample from these breeds. Karyotype of sheep, native pigs and of some native chicken
were also studied by the groups.
Genomic analysis has not been done for any species of domesticated livestocks.
Nepal is a good place for future genomic study of these domesticated animals.
Institutional

Structures
The Animal Breeding Division, Nepal Agricultural
Research Council isundertaking the leading role for studying the genetic resources and has the mandate forresearch has been given to NARC as an apex body. The Animal Breeding Division has
been working with Animal Breeding and Artificial
Insemination
Section of the
Department of Livestock Services.
The Institute of Agriculture and Animal Scienceteaches agriculture, veterinary and animal sciences.
Acknowledgments
The author would like to offer his sincere thank to Dr. Tsutomu Furukawa, Department
Genetic Resources; Dr. H. Seko, Genetic Resources Coordinator;
General;National

Instituteof Agrobiological

Resources,

Japan; for their invitation,encouragement and providing

to present

this paper.

The author is happy

Fisheries;

Mr. B.K. Baniya Director,

Executive Director,

Nepal Agricultural

Ministry

and Coordination

Research

Councilof Agriculture,

the financial

to acknowledge

Planningof Animal

and to Dr. Masahiro Nakagahra, Director
supportForestryand Fisheries,

and for organizing

the seminar

Dr. Upendra Mishra, Director,
and to Mr. Jagadishfor providing

LivestockandChandra Gautam theme the opportunity

to visit Japan topresent this paper.
References
Shrestha, N.P. 1994 Animal Genetic

Pradhan,Resources of Nepal and Their Conservation

Proceedingsof the Third

Global Conference on Conservation of Domestic Animal Genetic Resources. Editedby Crawford,

R.D.; Lister,E.E. and Buckley

J.T. Presented

University,

Kingston,

Canada.

Ontario,by Rare Breed International

1-5 August Queen's

S.L. and Rasali, D.P. 1994 Indigenous Animal Genetic Resources of Nepal: Potential for Their
Balanced Use Across EcologicalZones. Proceedingsof the Third Global Conference on
Conservation of Domestic Animal Genetic Resources. Editedby Crawford, R.D.; Lister,E.E. and241

Epstein,

Buckley

J.T. Presented

Ontario,

Canada.

H. 1977.

Domestic

Kawamoto, Y., Shotake,at Rare Breed International

AnimalsT., Adachi,of the native asses in Nepal
genetical
Agriculture,
Keshary,

studies

Workshopof Nepal. Holmes and Meier Publishers

Inc. New York and London.
B. 1988.

CentralBureau of Statistics.

Nishida,T., Shotake,T., Hayashi,

and their genetic

on the native

domestic

Y. and Kattel,

differentiation

form horses.

in Nepal.

andPart I: FacultyofAnimalGenetic

Resourcesin Nepal. Proceedings

Resourcesin Asia and Oceania,of Agriculture,

Forestryand Fisheries,

1 993

Sample Census of Agriculture

Nationalof SABRAO

Tropical

Agricultural

Research

199 1/92.

Kathmandu,

Nepal.

Japan.

T., Yamamoto,Y., Amano, T., Namikawa,T., Maeda, Y., Tsunoda, K., Rajbhandary,on the Interrelationship
Report Published

Program in 1988
1989.

variability

In: Morphologicalanimal and their wild forms in Nepal.

H.B., Kajune, T., Hongo, O., Kurosawa, Y, Douge, K., 1988/1989.
Investigation

Geneticof Tokyo, pp. 65.

N.P. 1980

on Animal Genetic

Centre, MinistryKingston,A., Nishida,

The University

K.R. and Shrestha

1-5 August Queen's University,and 1989.

Morphological

Between Native Domestic Animals

by Grant-in

Aid for Monobusho International

Part I and Part II, Facultyof Agriculture,

and Geneticaland Their Wild Forms
Scientific

The University

Researchof Tokyo,

GENERAL DISCUSSION

Chairpersons
Y. Tanabe
I. Bodo245

Questionsand answers in General Discussion

Comments by Dr. Tanabe
C Iwant to mention three topics that were not discussed in depth during the workshop.
1. More attention should be paid to the quality of products in animal production. Native
animal products are usually superior than those of modern breeds.
2. Adoption of modern engineering techniques to analyse meat quality without killingthe animal would be useful. For example, magnetic resonance imaging withcomputer tomography are useful techniques to detect the marbeling pattern of beef
cattle.
3. The mechanism ofheterosis should be elucidated. In maize, a high (0.8%) coefficientof correlation was observed between the yield (an expression of heterosis)
and thegenetic distance detected by DNA polymorphism (RFLP). No research has been
conducted on this point in animals. Genetic distance among strains or lines is easily
detected by randomly amplified polymorphic DNA. This technique will be very
useful to predict the expression of heterosis is chickens. (Tanabe)
Comments by Dr. Bodo
C Heterosis which can occur with preserved animal breeds should be mentioned.
Heterosis occurs when well adapted females (from ancient breeds) are crossed with
top males. (Bodo)
C Quality is a very difficult topic and it is difficult to agree on a definition because
different markets have different quality requirements. Thus, high quality is only thatwhich commandsa high market price. (Bodo)
Comment by Dr. Sekikawa
C. Disease resistance is important. In the next stage of conservation of animal genetic
resources the linkages between the phenotype and genotype (DNA polymorphic
markers) is necessary because both phenotypic and genotypic diversity are important.
( Sekikawa)
Comment by Dr. Ishii
C. Diversity is the key to conservationof animal genetic resources. I think selection

of

246

genetic resources to conserve on the basis of commercial quality is unwise, because
the standards fQr quality varies around the world as pointed out by Dr. Bodo. Peoples
perception of good quality also changes over time. So conservation of broad diversity
is most important. For preservation, making genomic libraries is most economic.
(Ishii)Comment by Dr Li Ning
C Wecan consider three levels of conserving animal genetic resources - animal level,
cell level and gene level. The methods and financial requirements for each differs.
There are also advantages and disadvantages
to each level. Of these three levels,
genomic libraries represent a powerful and efficient opportunity
for conserving
animal genetic resources. The genomic library can also be readily distributedformanyuses. (Li Ning)CLOSING

REMARKS249

Closing

remarks

HIDEFUMI SEKO
Genetic Resources Coordinator,
Kannondai 2-1 -2, Tukuba, Ibaraki

National

Instituteof Agrobiological

Resources

305, Japan

Thank you very muchDr. Tanabe and Dr. Bodo for your excellant chairmenshipof the general discussion.
Wehave nowreached the end of the indoor part of this workshop. I am sure you
are looking forward to seeing sunlight tomorrow during the outdoor part of the workshop.
On behalf of the organizing committee of the workshop, I should deliver a few
words.
Dr. Bodo reminded us that world wide conservation of native breeds andlandraces of animals related to agriculture is at a relatively early stage of development
compared with plants and microorganisms. But during this workshop a large amount ofdata and impressive results frommanyyears of research have been presented. It appears,
however, that in many countries much moreeffort needs to be directed towards animal
genetic resources conservation. We are all too well aware of the extinction and imminent
extinction of many animals. We cannot be reminded too often that "extinction isforever".
To effectively and efficiently conserve animal genetic resources many questions
have to be answered. As the discussion yesterday on the cryopreservationof chicken
semenindicated, we need to understand more about effective cryogenic preservation ofdelicate reproductive tissues.
I am very concious that in Asia meat consumption is increasing rapidly as people
across the region have greater spending power. As Dr. Matias discussed, in thePhilippines,
rapid economic development is resulting in a rapid increase in demand formeat. We are likely to be challenged in the future, to supply the demand for meat, much
morethan for the supply of agricultural
plant products for human consumption. It will
require concerted international
efforts to conserve animal genetic resources that will
provide the raw materials for improvement of farm animals. A vital task. However, meatproduction must be done in an environmentally sound way that does not threaten otherelements of the environment.

250

The MAFF International
Workshop on Genetic Resources aims at promoting
research exchange and collaborationon the development of technologies
and global
strategies
for conservation and use of genetic resources in national programs andinternational
research institutions.
This workshop has been of a small scale, but I
anticipate
that by continuing these efforts in the future, we will contribute to thepromotion of research and collaborationin conserving and evaluating animal genetic
resources.
As a plant breeder I have learned much over the last two days of intensive
discussions.
I thank everyone for their wholehearted participation.
I thank my colleagues here in Japan for their hard work in preparing papers forthis meeting.
I would like thank all the cooperating
institutes
in Japan and theAgriculture, Forestry and Fisheries Reseach Council for their support.
To friends who have come here from different parts of Asia and Europe it has
been marvelous to have you with us. We look forward to collaborationand interactionin the years to come. Wewish you a safe journey home when you leave at the end of theweek.
It is now my honor to close the indoor part of this workshop.
To all Happy Christmas and Happy New Year.
Thank you.

LIST OF PARTICIPANTS

The 3rd MAFF International
Workshop on Genetic Resources , December 5-7, 1995, Tsukuba,"Animal Genetic Resources : Efficient
Conservationand Effective Use

Japan

to

254

ABE, Tetsuya
Director, Institute of Radiation Breeding, NIAR, MAFF, JapanABLES, Gene Penaflor
Univ. of the Philippines-Los
Banos, Philippines
AMANO, Kazuhiro (Secretary
Staff)
Dept. of Breeding and Genetics, NIAI, MAFF, JapanAMANO, Takashi (Organizing
Committee)
Faculty of Agriculture, Tokyo Univ. of Agriculture,Japan
ANDO, Yasuo (Secretary
Staff)
Dept. of Animal Production and Grassland, JIRCAS, MAFF, Japan
ANH, Nguyen Quynh
Center for Transferring Science Technology, Nghe An, Vietnam
AOKI, Takayuki(Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, Japan
AOKI, Toshimitsu(Secretary
Staff)
Liaison and Coordination Division, AFFRCS, MAFF, Japan
ARAI, Tadao
Ibaraki Prefectural Institute of Swine Industry, Japan
ASTUTI, Maria (Session s)
Fukultas Peternakan, Universitas Gadj ah Mada, Indonesia
BODO, Imre (Keynote Address)
Dept. of Animal Husbandry, Univ. of Veterinary Science, Hungary
BOONGIRD, Somnuk
Entomology and Zoology Division Dept. of Agriculture,ThailandCHEONG, Ii Cheong.
National Livestock Research Institute, Korea
CHIBANA, Takashi (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, JapanDATTA, Rajit Kumar (Session 1)
Central Sericultural
Research and Training Institute, Central Silk Board,
India
DUVJIR, Suvd
(Tsukuba Univ., Japan), Mongolia
EBANA, Kaoru (Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, Japan
EBASHI, Shinji(Secretary
Staff)
Administration
Dept. , NIAR, MAFF, JapanFUKUI, Emiko
Faculty of Agriculture, Utsunomiya Univ. , Japan
FUKUTA, Katsuhiro(Session 5, Secretary
Staff)

255

Bio Defense Research Division, NIAH, MAFF, JapanFURUKAWA, Tsutomu (Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, Japan
GOTO, Nobuo
formerly Faculty of Agriculture, Kobe Univ. , Japan
HANADA, Hirofumi (Secretary
Staff)
Dept. of Breeding and Genetics, NIAI, MAFF, Japan
HANH Nguyen
Economic Department, Nghe An Province Committee, Vietnam
HARA, Wajiro (Secretary
Staff)
Dept. of Research Planning and Coordination, NISES, MAFF, Japan
HAYAKAWA,Hirofumi (Organizing
Committee)
Director, Dept. of Animal Production and Grassland, JIRCAS, MAFF, Japan
HAYASHI, Hiromitsu
School of Veterinary Medicine, Azabu Univ., JapanHIGUCHI, Mikito
Dept. of Animal Production, Tohoku National Agricultural
Experiment
Station, Japan
HINOMOTO, Norihide
Dept. of Insect Genetics and Breeding, NISES, MAFF, Japan
HORITA, Mitsuo (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, JapanHOSODA, Toshiaki(Welcome Address)
Research Councillor, AFFRCS, MAFF, Japan
IKEDA, Hidetoshi
Bio Defense Research Division, NIAH, MAFF, JapanINOUE, Hajime (Organizing
Committee)
Director, Dept. of Insect Genetics and Breeding, NISES, MAFF, JapanISHII, Susumu (Session 4)
Dept. of Biology, Waseda Univ., JapanISHIKAWA, Masaya (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, Japan
ITO, Hiroshi(Organizing
Committee)
Technical Director, NLBC, MAFF, Japan
ITO, Shin!ichi
Faculty of Agriculture,
Gifu Univ., JapanIZAIKE, Yoshiaki(Session 3, Secretary Staff)
Dept. of Reproduction, NIAI, MAFF, JapanKAGAMI, Hiroshi
Dept. of Breeding and Genetics, NIAI, MAFF, Japan

256

KAKIZAWA, Ryozo (Session 2)
The Yamashina Institute for Ornithology,
Japan
KAMIYA, Shin-ichiro
School of Veterinary Medicine, Azabu Univ., JapanKANAI, Yukio (Organizing
Committee)
College of Agrobiological
Resources, Univ. of Tsukuba, Japan
KANAZAWA,Takuya
Faculty of Agriculture,
Ibaraki Univ. , JapanKARIYA, Takayoshi
Director, Dept. of Reproduction, NIAI, MAFF, JapanKASAMO, Kunihiro
Director, Dept. of Applied Physiology,
NIAR, MAFF, JapanKASHIWABARA, Takao
Prof. Emeritus, Ibaraki Univ., Japan
KATO, Kunihiko (Organizing
Committee)
Director, Dept. of Genetic Resources I, NIAR, MAFF, JapanKATSURA, Naok i (Opening Address, Organizing
Committee)
Director, Dept. of Research Planning and Coordination, NIAR, MAFF, JapanKAWABATA,Shutaro
Society for Techno-innovation of Agriculture, Forestry and Fisheries, JapanKAWASAKI, Kenj iro
Dept. of Insect Genetics and Breeding, NISES, MAFF, JapanKIKUCHI, Kazuhiro (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, Japan
KOBAYASHI, Eiji
Society for Techno-innovation of Agriculture, Forestry and Fisheries, Japan
KOJIMA, Mieko
Faculty of Agriculture, Utsunomiya Univ. , Japan
KOMATSU, Masanori
Dept. of Breeding and Genetics, NIAI, MAFF, Japan
KOMIYAMA, Tetsuro
Shirakawa Institute of Animal Genetics, Japan
KOSEGAWA, Eiichi (Session 3)
Dept. of Insect Genetics and Breeding, NISES, MAFF, JapanKOSHIMOTO, Chihiro
Institute of Animal Serology, Tokyo Univ. of Agriculture,Japan
KOSUGIYAMA, Motoaki
Faculty of Agriculture,
Ibaraki Univ. , Japan
KUDO, Hiroshi
Dept. of Physiology,
NIAI, MAFF, Japan

257

KUNIHIRO, Yasushi (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, Japan
KUROSAWA, Yaetsu (Session 2)
Institute of Animal Serology, Tokyo Univ. of Agriculture,Japan
LI,Ning
Beijing Agricultural
Univ. , China
LI, PeiChang
Huangyuan Animal Husbandry College, Qinghai Province, P.R. of China,
China
LI, Rong Feng
The Research Center for Laboratory Animal Science, Inner Mongolia Univ. ,China
LY, Le Viet (Session 3)
Vice Director, National Institute of Animal Husbandry, Vietnam
MAEDA, Miki (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, Japan
MATIAS, Job Manaet (Session 3)
Dairy Training and Research Institute, Univ. of the Philippines
Los Banos,
Philipp ine s
MATSUKAWA, Tadashi
Director General, NIAI, MAFF, JapanMIKAMI, Hitoshi(Organizing
Committee)
Director, Dept. of Breeding and Genetics, NIAI, MAFF, JapanMINOBE, Yuzo
Director, Dept. of Molecular Biology, NIAR, MA?FF, JapanMIYASHITA, Susumu (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, JapanMORIYAMA, Hiromitsu
Dept. of Research Planning and Coordination, NIAI, MAFF, Japan
MULADNO
Faculty of Animal Science, Bogor Agricultural
Univ. , Indonesia
MURAMATSU, Susum (Organizing
Committee)
Faculty of Agriculture,
Utsunomiya Univ. , JapanNAGAI, Takashi
Dept. of Animal Production, Tohoku National Agricultural
Experiment
Station, Japan
NAGAMINE, Tsukasa (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, JapanNAITO, Mitsuru (Secretary
Staff)
Bio Defense Research Division, NIAH, MAFF, Japan

258

NAKAGAHRA, Masahiro
Director General, NIAR, MAFF, Japan
NAKAYAMA,Hiroki (Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, Japan
NAKAYAMA,Osamu
Faculty of Agriculture,
Ibaraki Univ., Japan
NAKAYAMA,Shigeki(Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, JapanNAMIKAWA, Takao (Session 2)
School of Agricultural
Sciences, Nagoya Univ., JapanNIRASAWA, Keijiro(Session 1, Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, JapanNISHIBORI,
Masahide
Faculty of Applied Biological
Science, Hiroshima Univ. , JapanNISHIKAWA, Tomotaro (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, JapanNISHIMURA, Marie (Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, JapanNOGUCHI, Junko (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, JapanNOMURA, Koh
Faculty of Agriculture,
Tokyo Univ. of Agriculture,Japan
NOZAWA, Ken
Faculty of Liberal Arts, Chukyo Univ., Japan
OBATA, Taro
Dept. of Research Planning and Coordination, NIAI, MAFF, JapanOCHIAI, Hirokazu (Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, Japan
ODA, Sen-ichi
School of Agricultural
Sciences, Nagoya Univ. , Japan
OGAWA,Kazutaka (Organizing
Committee)
Chief, Liaison and Coordination
Division, AFFRCS, MAFF, JapanOHSAWA, Katsuji
Director, Dept. of Cell Biology, NIAR, MAFF, JapanOISHI, Takao (Organizing
Committee, Secretary Staff)
Director, Dept. of Genetic Resources II, NIAR, MAFF, Japan
OKABAYASHI, Hisato
School of Veterinary Medicine, Azabu Univ. , Japan
ONIKI, Masaomi (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, Japan

259

ONISHI,

Akira (Session 4)
Dept. of Breeding and Genetics, NIAI, MAFF, Japan
OOI, Hideo
Institute of Silkworm Genetics and Breeding, JapanOYANAGI, Chie (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, Japan
SAKA, Norie
College of Agrobiological
Resources, Univ. of Tsukuba, JapanSAKAGUCHI, Minoru
Dept. of Animal Production, Tohoku National Agricultural
Experiment
Station, Japan
SAKAMOTO, Yuko
Azabu Univ., Japan
SANO, Akiko
Shotoku Women's Junior College, Japan
SASAKI, Sachiko (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, Japan
SATO, Kyoko (Secretary
Staff)
Liaison and Coordination Division, AFFRCS, MAFF, Japan
SEKI, Masumi (Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, JapanSEKIKAWA, Kenji (Organizing
Committee)
Director, Bio Defense Research Division, NIAH, MAFF, JapanSEKIYA, Mario
Faculty of Agriculture,
Utsunomiya Univ. , Japan
SEKO, Hidefumi (Organizing
Committee)
Genetic Resources Coordinator, NIAR, MAFF, JapanSHIMIZU, Manabu
Dept. of Reproduction, NIAI, MAFF, JapanSHIINA, Tsugio (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, JapanSHIOYA, Yasuo (Secretary
Staff)
Dept. of Reproduction, NIAI, MAFF, JapanSHIRATA, Kazuto (Secretary
Staff)
Dept. of Research Planning and Coordination,
NIAR, MAFF, JapanSRI Sulandari
BiologicalResearch Centre, Indonesian Institute of Science, Indonesia
SUTOPO
Faculty of Animal Science, Diponegoro Univ., Indonesia
TAGAMI, Takahiro(Secretary
Staff)

260

Dept. of Breeding and Genetics, NIAI, MAFF, Japan
TAKAGI, Nobuki
Utsunomiya Univ. , Japan
TAKAHASHI, Hideaki (Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, Japan
TAKEBE, Akira
Dept. of Breeding and Genetics, NIAI, MAFF, Japan
TAKEDA, Hisato (Session 4)
Dept. of Breeding and Genetics, NIAI, MAFF, Japan
TAKEDA, Kumiko
Dept. of Breeding and Genetics, NIAI, MAFF, Japan
TAMURA, Toshiki
Dept. of Insect Genetics and Breeding, NISES, MAFF, Japan
TAN, Soon Guan (Session 2)
Dept. of Biology, Univ. Pertanian Malaysia, Malaysia
TANABE, Yuichi (Keynote Address)
School of Veterinary Medicine, Azabu Univ., JapanTANAKA, Yoshio (Secretary
Staff)
Administration
Dept. , NIAR, MAFF, JapanTANIWAKI, Osamu
School of Veterinary Medicine, Azabu Univ. , JapanTOGASHI, Kenji
Dept. of Breeding and Genetics, NIAI, MAFF, JapanTOMOOKA, Norihiko (Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, JapanTSUGE, Michiyo (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, Japan
TSUKADA, Masami (Secretary
Staff)
Liaison and Coordination Division, AFFRCS, MAFF, Japan
TSUCHIYA, Kenichi (Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, JapanTSUTSUMI, Hisanobu
Utsunomiya Univ. , JapanUGAWA, Yoshihiro(Secretary
Staff)
Dept. of Genetic Resources II, NIAR, MAFF, Japan
UMEHARA, Masamichi(Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, Japan
VAUGHAN, Duncan (Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, Japan
WANG, Aiguo (Session
1)

261

College of Animal Science and Technology, Beijing Agricultural Univ.,
China
WANG,Wei-Hua
Dept. of Veterinary Medicine, Gansu Agricultural Univ. , China
YAGI, Takashi
Okayama Univ., JapanYAMADA,Yukio
formerly Faculty of Agriculture, Kyoto Univ. , JapanYAMADA,Yutaka (Session 5)
Dept. of Animal Production, Hokkaido National Agricultural Experimental
Station, MAFF, JapanYAMAMOTO, Akio (Secretary
Staff)
Liaison and Coordination Division, AFFRCS, MAFF, JapanYAMAMOTO, Yoshio
Faculty of Applied Biological
Science, Hiroshima Univ., JapanYAMANE, Akihiro
Global Environmental Research Division, National Institute forEnvironmental Studies, Japan
YOKOYAMA,Hidenori
School of Veterinary Medicine, Azabu Univ. , Japan
YOKOYAMA, Tadashi (Secretary
Staff)
Dept. of Genetic Resources I, NIAR, MAFF, JapanYUKUHIRO, Kenj i
Dept. of Insect Genetics and Breeding, NISES, MAFF, JapanZHANCHIV, Tsesrenjav(Session 1)
Institute of General and Experimental Biology, Mongolian Academy of
Sciences, Mongolia
MAFF: Ministry of Agriculture,
Forestry and Fisheries
AFFRCS: Agriculture,
Forestry and Fisheries Research Council Secretariat
NIAR: National Institute of Agrobiological
Resources
NIAI: National Institute of Animal Industry
NISES: National Institute of Sericulturaland Entomological
Science
NIAH: National Institute of Animal Health
JIRCAS: Japan International
Research Center for Agricultural
Sciences

Editors
Editor in chief
Managing editorsC

onsultingeditors

Oishi, Takao
Vaughan, Duncan A.
Furukawa, Tsutomu
Kikuchi, Kazuhii oShirata, Kazuto
Seko, Hidefumi
Kato, KunihikoPublishedby•EyR e s e r c h C o u n c il S e c r e ta r i a t o f M A F F a n d
N a t io n a l I n s titu t e o f A g r o b i o lo g ic a l R e s o u r c e s
K an n o n d a i 2 ‑1 ‑2 , T su k u b a , Ib ara k i 3 0 5 , Jap a n
I S B N 4 ‑9 9 0 0 1 1 0 ‑ 7 ‑4

M arc h , 1 9 9 7