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Biological consequences of Chernobyl:
20 years onAnders Pape Møller1 and Timothy A. Mousseau2
1

Laboratoire de Parasitologie Evolutive, CNRS UMR 7103, Université Pierre et Marie Curie, Bât. A, 7ème étage, 7 quai St. Bernard,
Case 237, F-75252 Paris Cedex 05, France
2
Department of Biological Sciences, University of South Carolina, Columbia, SC 29208, USA

The disaster at the Chernobyl nuclear power plant in1986 released 80 petabecquerel of radioactive caesium,
strontium, plutonium and other radioactive isotopes
into the atmosphere, polluting 200 000 km2 of land inEurope. As we discuss here, several studies have sinceshown associations between high and low levels ofradiation and the abundance, distribution, life history
and mutation rates of plants and animals. However, thisresearch is the consequence of investment by a few
individuals rather than a concerted research effort by theinternational community, despite the fact that theeffects of the disaster are continent-wide. A coordinatedinternational research effort is therefore needed tofurther investigate the effects of the disaster, knowledge
that could be beneficial if there are further nuclear
accidents, including the threat of a ‘dirty bomb’.

Introduction
One of the four nuclear reactors of the Chernobyl nuclear
power plant exploded on 26 April 1986 as a consequence ofhuman errors owing to a temporary shutdown of thecooling system. The explosion transported vast amounts ofradioactive material into the atmosphere, much of which
was subsequently deposited not only in the immediate
vicinity of the power plant in Ukraine, Russia andBelarus, but also over large parts of Europe and othercontinents (Figure 1). On this, the 20th anniversary of theworst environmental nuclear disaster in history, there is
still much disagreement among government agencies,
health professionals and scientists over the long-term
effects of low-level nuclear contaminants. The official UN
position [1] suggests that the consequences to humanhealth are much lower than expected, the park-likeappearance of the 2044.4 km2 Chernobyl exclusion zone,
with large mammals appearing to be increasing in
numbers, suggests an ecosystem on the rebound. However, the UN report, and interpretations of it in the
popular and scientific press (e.g. [2,3]), has generated an
optimism that might be unfounded.
Here, we discuss the information available concerning
the effects of the Chernobyl disaster on wild plant andanimal populations, and also humans. It is our hope that
this information will serve as a foundation for future
Corresponding author: Møller, A.P. (amoller@snv.jussieu.fr).

studies investigating the long-term ecological and evolutionary repercussions of chronic exposure to low-level
radioactive contaminants (low-level radiation has been
defined as the dose below which it is not possible to detect
adverse health effects, typically 1–20 rads). Given that theeffects of the disaster were felt on a continent-wide scale
but that research has generally been the result ofinvestment by a few individuals, we also call for acoordinated international research effort to further
investigate the environmental outcomes of the disaster.

A brief history of the Chernobyl event
On 26 April 1986, during a test of the ability of theChernobyl nuclear power plant to generate power while
undergoing an unplanned shutdown, safety systems wereturned off, leading to an explosion and nuclear fire thatburned for ten days, releasing between 9.35!103 petabecquerel (PBq) and 1.25!104 PBq of radionuclides into
the atmosphere (by contrast, the Three Mile accident in
Pennsylvania, USA on 27 March 1979 released just 0.5
terabecquerel). Although many of these radionuclides
either dissipated or decayed within days (e.g. 131Iodine),
137
Caesium (137Cs) still persists in the environment even
hundreds of kilometres from Chernobyl. Likewise,
90
Strontium (90Sr) and 239Plutonium (239Pu) isotopes arecommon within the exclusion zone. Given the 30, 29 and
24 000 yr half-lives of 137Cs, 90Sr and 239Pu, respectively,
these contaminants are likely to be of significance formany years to come.

Physiological and genetic effects of radiation
Immediately following the accident at Chernobyl, humans
exposed to high-level radiation suffered from acuteradiation sickness, including dizziness, vomiting andfatigue [1]. The long-term physiological effects of immediate and later exposure have also shown changes in levelsof antioxidants, immunity and hormones. Most of the longterm consequences of the Chernobyl disaster stem fromthe inhalation and ingestion of radionuclides generated by
the explosion and nuclear fire (Box 1), in contrast to theeffects that result from direct exposure to radiation from anuclear blast. The genetic consequences of radiation
exposure will depend on the received dose, dose rate andother indirect effects (Box 1).

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Figure 1. Distribution of radiation in Europe in May 1986 as a result of the Chernobyl disaster. Black squares indicate capital city locations. Adapted with permission from [31].

Antioxidant, immunity and hormone effects
Radiation can reduce levels of antioxidants, such as
carotenoids and vitamins A and E, which are used forprotecting DNA and other molecules from damage causedby free radicals [4–8]. Consistent with this prediction is
the finding that barn swallows Hirundo rustica studied
during 2000 from the most radiation-contaminated areas
had significantly reduced amounts of carotenoids andvitamins A and E in blood and liver, compared with birds
from control areas [9] (Box 2), although the level ofradiation was low compared with previous studies ofhumans [4–8] (Box 3). These reductions were the best
predictor of increased frequency of abnormal sperm frommale barn swallows in such areas [9].
Antioxidants can have important consequences for
immunity owing to their immunostimulating effects(reviewed in [10]). Studies of Chernobyl staff involved incleaning-up immediately after the accident revealed
impaired immune function compared with that of matched
control individuals [11–13]. Likewise, barn swallows from
Chernobyl had depressed levels of several types ofwww.sciencedirect.com

leukocytes (e.g. heterophils and lymphocytes) andimmunoglobulins, and reduced spleen mass comparedwith individuals from control areas [14], suggesting a
reduced ability to raise an efficient immune response. The
heterophil:lymphocyte ratio, which is an important
physiological indicator of stress [15], was also elevated inbarn swallows from Chernobyl compared with
control individuals.
The elevated frequency of partial albinism in barnswallows, humans and other organisms from the Chernobyl region can also be linked to an antioxidant deficiency[16,17]. Normal plumage or skin colour is produced by the
migration of melanocytes from the skin (so-called ‘melanoblasts’) to feathers; such migration can be disrupted ormelanocytes can die prematurely owing to low concentrations of antioxidants in the skin, resulting in albinism.
Mutations in genes encoding plumage colour can have asimilar effect [17], as shown by the finding that featherswith melanin-based colour are paler in barn swallows
from Chernobyl compared with those from individuals
from control areas [14]. Surprisingly, to our knowledge,

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Box 1. Radiation exposure, pathways and effects in animals
Pathways of exposure
There are several ways in which animals can be exposed to radiation. (i)Exposure by inhalation occurs as a result of breathing radioactive dust,
smoke or gaseous radionuclides into the lungs, where radioactive
particles often remain for a prolonged period*. Such exposure is most
relevant for a or b particle-emitting radionuclides (e.g. 239Pu or 137Cs
and 90Sr, respectively) because of the possible prolonged exposure to
the respiratory system. This is likely to be a major pathway of exposurefor those animals affected by Chernobyl fallout. (ii) Exposure byingestion occurs as a result of swallowing radionuclides. a and bemitters are again of greatest relevance owing to possible prolonged
contact with the digestive system. Also, because Sr and Pu are readily
absorbed, internal organs and tissues are also at risk; Sr and Pu are
more likely to be fixed in bones, teeth or liver, thus affecting
surrounding tissues for the life of the animal*. (iii) Direct or external
exposure is of most relevance for g radiation emitters (e.g. 137Cs) but oflimited relevance for a emitters because a particles cannot penetrate
the outer layer of skin. Contact with b emitters can also generate burns
or eye damage, but this requires close contact as b particles can travelfor only limited distances in the air*. Exposure by absorption andexternal exposure are of importance for plants and fungi.

Dose, rate and bystander effects
In all organisms, intracellular DNA is sensitive to radiation exposure
and is easily broken by exposure to low amounts of radiation. A
double-strand break is often lethal as it is difficult to repair correctly,
often leading to the loss of chromosomal material at cell division and,

there has been no study of the level of corticosterone orheat shock proteins from organisms from the Chernobyl
region, despite these being commonly used indicators ofstress in animals.
Mutation effects
In Table 1 we list 33 studies that investigated mutations or
cytogenetic effects of increased radiation around Chernobyl compared with control areas in a variety of plant andanimal species. This list is unlikely to be exhaustive, giventhat there are likely to be further studies published in
Russian, Belarusian or Ukrainian journals (usually onlyin the Russian language) that are thus relatively
inaccessible to western scientists. There is considerable
heterogeneity in the results, with 25 of the studies
showing an increase in mutations or cytogenetic abnormalities. Several studies showed an increase in mutation
rates for some loci, but not for others [18,19]. However,many studies were based on small sample sizes, with aresulting low statistical power being unable to show
differences of 25% as being statistically significant.
Only four of these studies investigated germline
mutations [18–21] and these all found significant
increases. Many of these studies were not included in
the review by the UN Chernobyl Forum Expert Group [1],implying that the conclusions of this group, that germline
mutational effects were weak if at all detectable, were notbased on available information.
Fitness consequences of the increases in mutation rates
or chromosomal aberrations remain largely unknown.
Ellegren et al. [19] reported an association between partial
albinism and reduced survival in barn swallows; asubsequent study of standardised differences in phenotype for over 30 different characters of barn swallows
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ultimately, cell death. Incorrectly repaired DNA can lead to mutations
and carcinogenesis [61].
Point mutations (i.e. single base-pair substitutions) generally show
a linear radiation dose response and are more prevalent at low doses.
Intermediate doses are often associated with frame shifts (i.e. single
base-pair insertions or deletions), whereas high doses often lead tomultiple mutations, which can generate intergenic lesions that resultin the loss of multiple genes. Such lesions increase with the square ofradiation dose [62].
The rate at which a radiation dose is received also influences
mutagenesis. In general, there is a linear relationship between
mutation rate and dose rate at low dose rates, and an exponential
response at high dose rates. Based on observations of DNA
repair-deficient cell cultures, low dose rates enable DNA repair,leading to lower mutation rates. Some intragenic lesions (i.e.
deletions) are induced by two-hit events (because DNA strands haveto be broken twice) and show dose-rate dependence [62]. Independent
multilocus mutations are sometimes generated by the same lowenergy track as a consequence of the folding patterns of DNA [62].
Recent studies of cell cultures exposed to highly focused low-doseradiation have shown so-called ‘bystander’ effects, whereby nontarget cells next to exposed cells show mutagenic effects. The
mechanisms underlying bystander effects appear to be diverse andreflect complex pathways of biochemical signalling amongcells [61].
* USEPA (2005) Exposure Pathways (http://www.epa.gov/radiation/
understand/pathways.htm).

between study populations near Chernobyl and in
relatively uncontaminated control areas revealed apositive relationship between difference in phenotype
and effect of the trait on mating success [22]. Mutations
with slightly negative fitness effects could easily be
exported out of the contaminated areas via organism
migration, with consequences for populations that havenot been directly exposed to radiation from the disaster.
Furthermore, accumulation of mutations in individuals
living in contaminated areas could increase the susceptibility of individuals to adverse environmental conditions,because mutants generally will show low levels of stressresistance, although this remains to be tested experimentally. Although the official UN report provided estimates ofhuman deaths attributable to the Chernobyl incident in
the order of 10 000 [1], we consider these estimates to bepremature, given the current level of knowledge ofmutational impact on humans and other organisms.
Therefore, more research is needed.
Ecological consequences of radiation
There has been little research on the ecologicalconsequences of the Chernobyl disaster, despite thefact that studies of the abundance of common speciesof vertebrates and invertebrates can be done easily andat a low cost.
Life-history effects
Life-history consequences of radiation are expected
because life-history traits are generally affected byphysiological pathways that, on their own, and incombination, can be affected by mutations or thephysiological effects of radiation on antioxidant levels[23]. Studies of the barn swallow have shown significant

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Box 2. Medical effects of radiation
Radiation emergency workers at the Chernobyl plant during and
immediately following the accident received very high doses ofradiation and, in total, 57 died. Subsequent investigations of the
medical effects of this radiation suggested increased rates ofcongenital defects, cancers and cardiovascular disease in exposed
humans compared with controls. For example, in contaminatedareas near Gomel and Mogilev, Belarus, thyroid cancer arising frominhalation of radioactive dust has increased significantly since 1986,
as have congenital defects and spontaneous abortions (Figure I).
This is in contrast to the finding of no increase in frequencies of suchconditions in several Western European countries over the sametime frame [63].
Several studies have also reported increases in the frequency ofthe medical effects of radiation in control areas, where humans were
not exposed to elevated radiation levels [1] (Figure I). This raises
serious problems of interpretation, with some scientists suggestingthat effects of ‘worry’ or improved reporting rather than radiation are
the cause of such increases. Alternatively, the transition from
communism to free market societies around 1990 caused reductionsin income, nutritional condition and medical services for much of thehuman population, which could also explain these results. Unlessthe confounding effects of such changes can be controlled
statistically, there is little possibility of interpreting available medical
records reliably. This situation also makes studies of animal or plantmodels all the more important, both because of their shorter
generation times and the lack of importance of ‘worries’ as a causeof any health effects.
The encouraged resettlement of contaminated zones for agriculture in Belarus is based mainly on predictions that !10 000 people
are likely to die of Chernobyl-related cancers. Even the best-casescenarios do not include non-cancer mortality or increased
morbidity of any sort. Neither is there any assessment of thehuman costs associated with medical treatment. Given the unprecedented nature of the Chernobyl disaster, it seems prudent to besensitive to the possible unpredicted impacts, given that many
human diseases have long latency periods (e.g. smoking-related
illnesses often only occur following 20–30 years of exposure).

(b)
8
20
Thyroid cancer rate

Frequency per 10 000 inhabitants

(a)

15

10

5

0
BeforeChernobyl
event

After
Chernobyl
event

6

4

2

0
1985

1990

1995
Year

2000

2005

Figure I. Radiation and human health consequences. (a) Increased frequency of
spontaneous abortion (red bars, !0.1) and congenital malformation (whitebars) in Gomel and Mogilev, Belarus before and after the Chernobyl disaster.
Mean values are shown (s.d.). (b) Increased frequency (per 10 000 inhabitants)of thyroid cancer in children in relation to radiation levels in Belarus (circles)
and Ukraine (squares) before and after the Chernobyl disaster, showing an
increase after the disaster. Adapted with permission from [1,64].

relationships between background level of radiation andthe timing of reproduction, clutch size and hatching
success [24]. Non-breeding females are generally uncommon in temperate-zone passerines, but 23% of female barnswallows from Chernobyl were non-breeders and lacked anaked brood patch during the breeding season (breeding
female birds moult feathers on their belly to facilitatewww.sciencedirect.com

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incubation of eggs), whereas this fraction was close to zeroin the control area in Ukraine [24]. The fraction of nonbreeders was negatively related to level of backgroundradiation at different sites [24]. Although breeding datehas a strong impact on the probability of recruitment inbirds [25], there was no delay in breeding associated withan increase in background radiation of two orders ofmagnitude [24]. The clutch size of barn swallows was
reduced in sites with elevated radiation, and hatching
failure was associated with background radiation level inthis species [24].
Adult survival prospect is an important determinant oflife-time reproductive success [26] and any reduction insurvival rate will have important fitness consequences.
Adult barn swallows breeding in Chernobyl had survival
rates that were reduced by 24% and 57% for males andfemales respectively, in comparison with control areas[24]. These differences are large compared with normal
intraspecific variation in survival rate.
Reduced adult survival and reproduction suggeststhat extant populations of these bird species in this area
are unlikely to be viable; only if there is significant
immigration from source populations to the Chernobyl
sink can these populations be maintained. Based on ourcurrent knowledge of source–sink dynamics [27] andknown sex differences in adult survival rate anddispersal rate in the barn swallow, we can predict thatthe rate of immigration should be considerably greater
into Chernobyl than into control areas, but only after
1986. Migratory birds winter in specific areas where theytend to return to in subsequent years, and thegeochemical fingerprint of these wintering grounds isstored in the stable isotope composition of feathers for
those species that moult during winter [28]. This fact
was used to investigate the extent to which variance instable isotope profile differed between barn swallows
from Chernobyl and control areas before (using museum
material; Box 3) and after the disaster [29]. Stable
isotope profiles from before and after the Chernobyl
disaster were more heterogeneous than were those of thecontrol population from control areas. Variances in stablecarbon isotope content of feathers (d13C) of both sexes
from post-1986 samples from Chernobyl were significantly larger than variances for feather samples from thecontrol region, and compared with variances for historical samples from both regions. This suggests that stable
isotope measurements provide information about population processes following environmental perturbations.
It also suggests that optimistic prospects for the future ofanimal and plant populations reported by the UN
Chernobyl Forum [1] are biased because apparently
healthy populations might be sink populations rather
than sources exporting individuals elsewhere.
We can only speculate about the underlying mechanisms that cause the effects of radiation on life history. One
possibility is that the reduction in body antioxidant levels
directly affects the timing of reproduction, clutch size andsurvival prospects because female reproduction is limited
by antioxidant availability [30]. Similarly, a reduction in
antioxidant levels associated with radiation might also
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Box 3. Problems of analyzing effects when there is only one event
The Chernobyl disaster provides an unrivalled opportunity to test theeffects of radiation on biological phenomena under large-scale field
conditions, given that it is not straightforward to extrapolate from thelaboratory to the field. However, this also raises philosophicalconsiderations about how to use a single observation to makerigorous tests of scientific hypotheses. We can imagine three different
solutions to this problem, of which two have been adopted so far.
First, investigations that simultaneously use temporal and spatial
patterns of a phenomenon can compare the situation before and after
the disaster in contaminated and control areas without radioactive
contamination both before and after the disaster. Such an approachhas been used to study partial albinism and asymmetry in feathers of
the barn swallow Hirundo rustica before and after the Chernobyl
disaster (Figure I; [65]). Partial albinism is caused by mutations that are
rare among animals, but the frequency in swallows has increased in
the Chernobyl region by five-tenfold since 1986, but not in control
areas (Figure I). Similarly, the degree of asymmetry in the length ofoutermost tail feathers of barn swallows has increased fivefold in
Chernobyl, but not in control areas [65]. For many species,
comparisons of samples before and after the disaster will not be
feasible because of the lack of material pre-dating the disaster.

(a) (i)(ii)

Second, whereas a given pattern might arise for random reasons
with a sample of two, a pattern that occurs repeatedly is unlikely toarise by chance. This approach was used for studies of bilateralasymmetry (e.g. differences in perfect symmetry between the lengthof right and left characters, such as right and left sides of leaves andright and left wings) in plants and animals in radioactively contaminated areas near Chernobyl and in control areas for a total of 15
species (four plants, four insects, two fish, one amphibian, one bird
and three mammals [66–68]). All revealed higher frequencies ofasymmetry in representatives from Chernobyl, deviating significantly
from the binomial null expectation [69]. However, this method
requires many studies of different species.
Third, the heterogeneous spatial distribution of radiation implies
that it is unlikely that a similar spatial distribution of phenotype
will occur by chance. Thus, an analysis of spatial autocorrelation
or a Mantel test is unlikely to provide a significant relationshipbetween radiation and phenotype unless there is an effect ofradiation on the phenotypic trait in question, especially if
geographical distance is controlled statistically. This approach
has yet to be used, although it will provide a useful alternative topreviously used methods.

(b)Fequency of partial albinism (%)

20

15

10

5

2005

2004

2003

2002

2001

2000

1996

1991

Before 1986

0

Year
Figure I. Partial albinism in barn swallows. (a) Barn swallows Hirundo rustica with (ai) and without (aii) partial albinism. (b) Frequency of partial albinism in barn swallows
Hirundo rustica in Chernobyl (red bars) and control areas (white bars) before and after the disaster. Adapted with permission from [70].

migratory birds that, during the annual migration,
produce large amounts of free radicals that must be
eliminated to avoid damage to DNA and othermolecules [10].
Future prospects for Chernobyl research
Chernobyl constitutes the most extensive ‘natural’ field
laboratory for studies of effects of radiation and researchduring the past 20 years has revealed important insights
into the consequences of low- and high-level radiation.
However, this ‘facility’ has yet to be fully exploited. It issurprising that there are only a few studies of mutation rates
in a small number of species, and that the ecological andevolutionary consequences of low-level radiation remain
poorly known. No study has, to our knowledge, investigatedwhether the disaster has had any effects on population
densities of common plants or animals. Likewise, no study
has, to our knowledge, attempted to determine whether
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slightly deleterious mutations arising from Chernobyl are
migrating out of the contaminated zone.
This lack of progress is likely to be a result of the
low level of investment in research at Chernobyl.
Consider the 11 September 2001 event in New York,
which resulted in OUS$100 billion in funding for all
kinds of research, including military research. By
contrast, the Chernobyl disaster has attracted !
US$10 million over the past 20 years. This lack offunding is far from what one might expect given the
non-negligible threat of a ‘dirty’ bomb, the use of
nuclear weapons and possible further accidents at
nuclear power plants. Even the nuclear power industry
and the over-seeing government agencies should have astrong interest in large-scale research to determine theeffects of radiation exposure.
We believe that a concerted research effort, funded by
the EU, USA, Japan and, to a lesser extent, local

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Table 1. Studies investigating the effects of radiation in Chernobyl on cytogenetics, genetic variability and mutations
Species
Chromosome aberrations
Human Homo sapiens

Yellow-necked mouse
Apodemus flavicollis
Mouse Mus musculus
Channel catfish Ictaluruspunctatus, Crucian carp
Carassius carassius, carp
Cyprinus carpio, tench
Tinca tinca
Oligochaetes: Dero obtuse,
Nais pseudobtusa, Nais
pardalis
Scots pine Pinus sylvestris
Somatic mutations
Human H. sapiens

Bank vole Clethrionomys
glareolus

Genetic marker

EffectComments

Refs

Lymphocytes withchromosomal aberrations

Increased by a factor 2–10

In women from Gomel andMogilev, Belarus
In clean-up workers
In children from Belarus[32]

Chromosomal aberrations
Number of reciprocaltranslocations
Frequency of aneuploidy

Fruit-fly Drosophilamelanogaster
Wheat Triticum sativum
Thale cress Arabidopsis
thaliana

Germline mutations
Human H. sapiens

Barn swallow Hirundo
rustica
Other effects
Mouse M. musculus
Wheat T. sativum, rye
Secale cereale[34,35]

Increased aneuploidy incontaminated areas[36]

In field populations[37]

Chromosomal aberrations

Increased by a factor 3

In field populations[38]

Minisatellites

Increased rate
No significant increase
No significant increase[39,40][41][42]

No significant increase
No significant increase

[43]
[44]

Multiple substitutions andtransversions wererestricted to samples from
Chernobyl
Increased by 19% in
mutations and by 5% inheteroplasmy, although
not significant
No significant increase[45]

Minisatellites andmicrosatellites
Microsatellites
Mutations

Point mutations

Mitochondrial Cytochrome bheteroplasmy
Breakage in DNA
DNA content based on flow
cytometrySex-linked recessive lethal
mutations
Microsatellites
Lethal mutations
Lethal mutations

Scots pine P. sylvestris

Increase by a factor of 15

Increased by a factor w2

Mutations and heteroplasmy

Channel catfish I. Punctatus
Crucian carp C. carassius

Mutation rate at enzyme lociProtein-coding genes
Minisatellites

RAPDs
Microsatellites

Lethality, embryo mortality andsterility
Aberrant cells

No significant increase[46]

Transplant experimentwith exposure during
90 days
Short-term transplant
experiment

Increased rate of breakage
Changes in DNA content,
but unrelated to known
measures of contamination
Increased
Increased by a factor 10
Increased by a factor 2–4
Rate 4–8 times higher thanin controls in 1992
Increased by a factor 20
Increased by a factor 4–17
Increased
Increased by a factor 1.6 inmen only
Increased rate
Increased by a factor 2–10

Increased
Increased in a dosedependent manner

governments, is needed. Such an effort could coordinate
research, establish a modern research facility and boost
local scientific competence. A major international instituteof radiation research supported by the internationalwww.sciencedirect.com

[33]
[33]
[33]

Chromosomal aberrations

Substitutions in cytochrome b

Mouse M. musculus

Increased by a factor 10–20
Increased by a factor 3–7
Increased by a factor 3–7[47–49]

[50]
[51]
[52]

[53]
Transplant experiment
In greenhouse and field
populations

[54]
[55]
[56]

In field populations
In field populations

[55]
[55]
[18]
[21]

Increased in only two outof three microsatellites
Outcomes of matedlaboratory animals[20,57]
[19]

[58]
[59]

community would make the most out of one of the largest
man-made environmental disasters, to the benefit of thelocal community, the general scientific community and theworld at large.

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Acknowledgements
We thank G. Milinevski, A. M. Peklo, E. Pysanets, N. Szczerbak, S.
Gashack, M. Bondarkov and A. Tokar for logistic help during our visits to
Ukraine. We received funding from the CNRS (France), the University ofSouth Carolina School of the Environment, Bill Murray and the SamuelFreeman Charitable Trust, The US Civilian Research and Development
Foundation, the National Science Foundation and National Geographic
Society to conduct this research.

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