CONE ANALYSIS OF

SOUTHERN PINES -*GUDEB~K
About the Authors

This Report was prepared jointly by the Southeastern Forest Experiment Station and the Southeastern Area, State and Private Forestry,

USDA Forest Service. Names, titles, and locations for Southeastern

Station authors are :

D. L. Bramlett, Principal Plant Physiologist, Macon, Georgia; G. L.

DeBarr and H. 0. Yates Ill, Principal Entomologists; T. Miller, Principal

Plant Pathologist; and K. D. Ware, Chief Mensurationist, all of Athens,

Georgia. _’

Southeastern Area authors are :

‘1

E W. Belcher, Jr, Director, and R. P. Karrfalt, Assistant Director,

Eastern Tree Seed Laboratory, Macon, Georgia; G. D. Hertel, Seed

Orchard Insect Program Coordinator, Pineville, Louisiana; an# C. W.

Lantz Nursery-Tree Improvement Specialist, Doraville, Georgia.
I.

II.

III.

IV.

V.

VI.

VII.

VIII.

IX.

X.

XI.

XII.

TABLE OF CONTENTS

INTRODUCTION

CONE AND SEED DEVELOPMENT AND MORPHOLO&

GENERAL GUIDELINES FOR CONE SELECTION

RECOGNIZING ABORTED OVULES AND SEEDS .’

CONE-ANALYSIS PROCEDURE

CAUSES OF OVULE AND SEED LOSSES

SUMMARIZING DATA

DATA INTERPRETATION AND UTILITY

FOR ADDITIONAL INFORMATION

GLOSSARY

LITERATURE CITED

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.4

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14

17

19

22

23

24

APPENDIX . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ...26
PREFACE

The rnformatron in thus publrcatron was presented ai’ two Cone Analysis

W orkshops organrzed by Dr Earl W Belcher, Jr, Director of t$ Eastern Tree Seed

Laboratory, durrng November 1976 at Macon, Georgia. These workshops were presented by personnel from the Southeastern Area, State and Pnvate Forestry. and the

Southeastern Forest Experiment Statron The authors of this workbook are those who

developed the workshop and conducted training sessrons.

Thus gurdebook contarns the rnformatron essential for cone analysis and rnterpretatron of the results Cone Analysrs Service (CAS) IS also available at cost from the Eastern

Tree Seed Laboratory at Macon, Georgia

2
CONE ANALYSIS OF SOUTHERN PINES

A Guidebook

D. L. Bramlett, E. W. Belcher, Jr., G. L. DeBarr,

G. D. Hertel, R. P. Kat-rfalt, C. W. Lantz, T. Miller,

K.D. Ware, and H. 0. Yates III

I. INTRODUCTION

Southern pine tree improvement programs require an ample supply of Improved seeds, 1 but productron from southern

pine seed orchards has often been disapporntrng If high productron IS to be malntarned. yields must be monitored and

causes of seed losses must be Identified. Techniques for determrntng seed efficiency were first used for red pine, Pinus

resinosa Art.. by Lyons (1956) Bramlett (1972b. 1974) modrfred and further developed a procedure. known as cone

analysis, for evaluating production efficrency In southern pine seed orchards

Cone analysis provides rnformation needed to evaluate seed productron and seed orchard management Actual seed

yield of rndividual cones is compared to the potentral seed yield Productrvrty can then be expressed In terms of seed efficiency. One can determrne in which stages of seed development certarn losses occur, and the types of seed failures can be

identified and quantrfred.

This guidebook outlrnes the basic cone-analysrs procedure, the factors Involved In losses of seed, and the rnterpretatron

and value of the results It is a stepwise gurde for those who wish to conduct their own cone analyses More detailed drscussions may be found in the publicatrons lrsted In the Literature Cited section Tree Improvement workers who do not

want to analyze cones themselves can obtarn this sen.lce at cost from the Eastern Tree Seed Laboratory This guidebook

will help them understand the analytical results they receive

tTerms in boldface type are defined in a glossary at the end of thus gurdebook

3
II. CONE AND SEED DEVELOPMENT

AND MORPHOLOGY

A basic knowledge of cone and seed development and morphology IS essential to understand cone analysis.

Pine trees produce two types of strobili: male or pollen-producing catkins and female or ovule-producing cones The

pnmordia of the female cones are initiated in the summer The followlng spring female flowers emerge In an erect POSItion from protective bud scales on the tips of new shoots (fig. 1A). The female flowers are composed of many soft, fleshy

scales spirally attached at right angles to a central axis. Pairs of ovules originate as small protuberances at the bases of

scales (fig. 1 B). Not all scales produce functional ovules; only those In the central region of the cone have the potential to

produce ovules and eventually seeds. These are called fertile scales. The lower scales and those at the tip of the cone are

infertile and never bear seeds (infertile scales).

At the time of pollination, a single layer of cells (integument) covers the ovule. Pollination occurs when the pollen

grains are carried by the wind from the male catkins to the female flowers. Pollen grains enter the ovule (fig. 1Cl through

an opening (the micropyle) In the seedcoat. Once in contact with the ovule. pollen grains germinate and pollen tubes

grow from the pollen chamber into the ovule tissue (nucellus) (fig. 1 D). The pollen tube develops slowly during the first

growing season, and the winter is over before it reaches the mature egg cell-bf the ovule. For more than 1 full year. the

ovule remains only a fraction of the size of a fully developed seed. During the year after pollination, the female flowers are

called conelets. :.

Gametophyte tissue

Ovule Fertilization

a, \X ’

P

‘, ;:: Nucellus , \F

' “~M~cropyle

Female Flower

J

-Male Catkin

Gametophyte tissue

H

Cotyledonsc$.

HypocotylA \ ,i

4

PRadicle

Seedling

Figure 1. -Pine seed formation. Drawing adapted from one provided by Carolina Biological Supply Company,

4
When vegetative growth starts In the second spring, the ovule also resumes development and enlarges. From 12 to 16

months after pollrnatron, depending on prne specres. the pollen tubes grow far enough into the ovule to reach specialrzed

structures (archegonia) contarning the egg cells (fig. 1 E). Sperm cells from the pollen tubes fertilrze the eggs, and embryos

(2N tissues) form. The gametophyte trssue surrounds the embryo and provides the nutritronal reserves for the seed (1 N

trssues). Dunng this time. the conelet enlarges rapidly to form a full-srzed cone. Seeds are full sized at the trme of

fertilization, but seeds and cones continue to mature until late summer or early fall

A mature, fully developed seed (fig. 1 G) has an embryo composed of Immature needles (cotyledons), immature stem

(hypocotyl), and an immature root (radicle). The gametophyte tissue enveloprng the embn/o provrdes the food reserves for

germination. The hardened seedcoat protects the seed

The sequence of pine seed development - from flower pnmordra formatron to seed fall - IS illustrated in figure 2 This

development spans parts of three growing seasons.

JIVCJ HI IHIIY 1 /

IYEAR 01 ‘\ \ \

Figure 2.-The cycle for flowering and seed production in southern pines.
III. GENERAL GUIDELINES

FOR CONE SELECTION

The method and number of cones to select for analysis depend on the analytrcal objectrves. the planting desrgn. ant

vanatron wrthrn the orchard

Overall performance of the orchard can be evaluated by selectrng cones from throughout the orchard. Exclude all rnsectdamaged cones, but do Include cones of all sizes that are normally harvested and extracted Such a bulk sample can bs

used for comparrsons with other orchards or with other years for the same orchard A sample of from 25 to 100 cones per

orchard IS normally suffrcrent to evaluate overall seed effrcrency However, wrth such selectron. no conclusrons can bs

reached about rndrvrdual clones.

If seed effrcrencres must be estimated for rndrvrdual clones, cones have to be collected separately from specrfrc clones

From lrmrted data, It appears that 10 or more cones per clone are required If rndrvrdual ramets are sampled wrthrn a clone

then three to five cones per ramet wtll probably be necessary Addrtronal research and experience will provrde a better basis

for specrfrc cone sample numbers

How well the sample represents orchard condrtrons depends on the vanatron wrthrn the orchard. Morphologrcal traits

such as cone length, number of scales, and seed potential vary relatively Irttle,‘and only small samples are required to estimate these traits wrth specified relrabrlrty. Seed-yield traits, however, are more‘idrffrcult to estimate because multiple factors (pollrnatron. Insects, etc.) contnbute to the varratron. Seed and ovule losses’ must be attnbuted to specrfrc agents tc

Identify problems In orchard management, and such losses can be highly vanable. Since seed losses are less wher

management IS good, seed-yield traits can be estimated from relatively small cone samples rn a high-yreldrng. wellmanaged orchard The very orchards where problems exist and where rnformatron needs are greatest are the most difficuli

to assess and require the largest cone samples

IV.: RECOGNIZING ABORTED

OVULES AND SEEDS

In order to understand the details of the cone analysrs procedure, It IS first necessary to be able to distinguish a

developed seed from an aborted ovule. Ovule abortion may occur during the frrst or second year of development. Frrstand second-year aborted ovules can be drstrnguished from developed seeds fairly easily by the methods grven in thus

section

FIRST-YEAR ABORTED OVULES

This class of ovule abortion occurs on fertile scales during the conelet stage - before cone enlargement begins in the

second growing season The seed wing develops normally, and these ovules have been called “wings without seeds”

Rudimentary, nonfunctronal ovules may develop with wings on some of the larger rnfertrle scales. Infertile scales can be

recognized by therr narrow bases (fig. 3). The nonfunctronal ovule never has the capacity of becoming a seed and appears

as a small dark spot at the base of the wing (Lyons 1956). First-year aborted ovules are noticeably larger than rudimentary

ovules (figs 4. 5A and 58). Some rnfertrle scales at the base of a cone may produce wings with no evidence of even a

rudimentary ovule. The smallest rnfertrle scales at the very base of the cone may not produce wings.

SECOND-YEAR ABORTED OVULES

Ovules that abort during the second growing season are always larger than those that abort during the first year They

may have a partrally developed seedcoat (figs 4B and 5C) These ovules are usually considerably smaller than developed

seeds, because they generally abort early In the second year of development Some ovules abort In late spring or early summer, these may be as large In outline as fully developed seeds but their seedcoats are collapsed. These ovules are frequently necrotic or shriveled and usually are attached to the cone scale with resin (fig. 50).

DEVELOPED SEEDS

Developed seeds are full sized and have complete seedcoats (ftgs. 3C and D; 4C: 5E. right. and F). These can be placed

in specrfrc classes based on thetr internal appearance on a radiograph (X-ray picture).

6
Figure 3.-Cone scales- (A & B) infertile scales with narrow base and no functional ovules, /C & DJ fertile scales containing two

developed seeds with mature seedcoat.

Figure 4.-Classification of aborted ovules and developed seeds: (Al first-year aborted ovules, (BJ second-year aborted ovules, and iC1

developed seeds.
Figure 5.-Classification of ovules and seeds: (A) rudimentary ovules from infertile scale; (BJ left. first-year aborted ovule; right normal

OVUle; (cl left early second-year aborted ovule; right normal ovule; (0) left, typical seedbug induced second-year aborted ovule; right

normal ovule; (El sliced seed: left empty seed: right filled seed; (F) fully developed seeds.

8
I/. CONE-ANALYSIS PROCEDURE

In cone analysis, all loose seeds and aborted ovules are extracted from the cone by normal procedures Then cone scales

re systematrcally removed and all the remarnrng ovules and seeds are collected The number of scales classed as fertrle

rg 3) IS used to determine the brologrcal potential of the cone to produce seeds Ovules and seeds extracted and drssected

om the cone are then classified by type (figs. 4 and 5) The seed efftcrency of each cone IS the number of filled seeds as a

ercentage of the brologrcal potential The causes for seed losses can be Identified and quantified on radiographs The ex-

!nt of seed losses to each cause indicates where correctrve measures mrght be taken to reduce seed mortality.

ClUlPMENT NEEDS

Cone analysrs requires ltttle specralrzed equtpment Essentral needs are.

1.

2

3

4

Paper collectron bags for individual cones (bag size depends on tree species) _.

Cone-drying facilrty (90°-1 05” F 132”.40” Cl and relative humrdrty below 50 percent)

Werghrng balance accurate to 1 gram :.

Variable-speed 3%Inch electnc drill mounted tn a stationary honzontal posrtron and adjusted to run at the slowest

speed

5 Drill bits of various sizes (inches). l/4, 5/l 6, 3/8, 7/l 6, l/2. 5/8.

6. Grafting knife

7 Leather work gloves

8 Metric ruler

9 Containers for soaking cones

10 Germrnatron suppltes and equipment

11 X-ray supplres and equipment

ROCEDURE

The stepwise procedure for cone analysis IS presented In a flow chart In figure 6 Bits of data that should be recorded are

drcated In the chart, the columns mentioned are those on the sample data sheet In the Appendix Some morphologrcal

arts are considered optronal (shaded columns on data sheet) and are not necessary for computatron of the most Important

?ed and cone charactenstrcs.

Two cone-analysrs procedures are presented These are designated “Procedure A” and “Procedure B ” During the

)Ilectron of data, there IS one basic difference In the two procedures With Procedure A, the extracted and dissected

:veloped seeds are combined (steps 9A and 15A ~ columns 46-61) With Procedure B, the extracted developed seeds

tep 9B - columns 46-61) are kept separate from the dissected developed seeds (step 15B ~ columns 62-77) from the

)ne. Each procedure has certain merits andlrmrtatrons. which are summarized below

Procedure A. ~ This procedure evaluates the yield and the extraction and germrnatron effrcrencres of all the seeds In the

ature cone The developed seeds extracted from the cone by normal procedures are combrned with the developed seeds

covered by removrng the cone scales (drssectron) All seeds are then radrographed and classrfred as described In sectron

The advantage of the procedure IS that only a srngle radiograph IS required for each cone Once the cone analysrs I S

:gun on a parttcular cone, the extractron. drssectron, and seed classrfrcatron can be completed at one time Also, the

rmary emphasis IS on the number of seeds actually produced In the cone, rather than just on those seeds extracted from

e cone under laboratory condrtrons. This procedure IS srmplest for small organrzatrons doing their own cone analysrs GerInation percentage using Procedure A IS based on the ratro of the number of germinated seeds to the total number of

led seeds produced by the cone.
Procedure B. ~ Thus IS the procedure currently berng used by the Eastern Tree Seed Laboratory for its Cone Analysrs Servrce (CAS) In this procedure the seeds extracted after forced drying are radiographed, and tests of their germination are

begun before the remarnrng steps rn the analysrs are completed. Because germination testing IS the step that requires the

most time. this procedure can substantrally reduce the number of weeks needed to complete the analysis. This saving IS

partrcularly valuable when many cones are to be analyzed.

The germlnatron percentage In Procedure B IS the percentage of the extracted filled seeds that produce a normal seedling. Extraction efficiency by this procedure IS the percentage of all filled seeds in the cone that were extracted after dryrns

(Karrfalt and Belcher 1977).

Before startrng cone analysrs. the user must decrde whrch procedure to follow. Once a procedure IS decrded upon, the

steps and computatrons Indicated for that procedure must be followed

Figure 6. -Flow chart for the steps in cone analysis. At several points steps are &ferent for Procedure A than for Procedure B.

Step 1

Collect a sample of mature, healthy cones to represent the seed

orchard or specific clones (see section III).

c

Step 2

Place each cone In a separate bag with species, year, orchard,

clone, ramet and cone identification.

(Record: columns 5-27) I

I Step 3

Air-dry cones until scale separation begins.

4

Step 4

Dry cones In forced draft oven or kiln at 90°-105” F (32O-40” C)

for 24 hours. -A

4

Step 5 (optional)

Estimate and record degree of cone opening. 1 = completely

closed; 2 = one-third open, 3 = two-thirds open; 4 = only slrghtly

closed: 5 = completely open (Record: column 28)

Step 6

Extract loose seeds and aborted ovules by banging cone on counter

top.

J

10
Step 7

Separate developed seeds from aborted ovules (see sectlon IV).

4

Step 8

Count and record number of extracted, developed seeds. (Record.

columns 43-45)

Place developed seeds In a labeled

b RadIograph seeds In envelope :‘.

c Begin germlnatlon procedures (see step 18)

d ClassIf) seed Images on radiograph (see sectlon VI) (Record columns 46-61)

t

Step 10

Separate first-year aborted ovules from second-year aborted ovules

(see sectlon IV) Record only number of second-year aborted

ovules (Record columns 41-42) Discard all aborted ovules

Step 13

Dissect cone beginning at base of cone, remove lower scales with

a knife point (fig 7) As cone axis IS exposed, use a drill with bit

diameters of l/4 Inch to 5/E Inch to remove cone axis This

facllltates scale removal Alternate dnlllng (1 Inch or less at a time)

and scale removal (fig 8) Match drill bit diameter with that of the

cone axis

1

+

11

,

Step 1 1

Soak cone In water until scales close

-c

Step 12 (optional)

Measure cone length to nearest mllllmeter (Record columns 29-

31)

I
Step 14

Separate scales into groups: (1) lower infertile scales, occurring at

the base of the cone (figs. 3A. B and 5A); (2) fertile scales, occurring in the central two-thirds of the cone and having two functional

ovules or developed.seeds (figs. 3C and D); and (3) upper infertile

scales, occurring at the apex of the cone and not having the

capacity to produce seed.

Step 15A

a. Take developed seeds from removed cone scales

and add to previously extracted seeds in the envelope (step 9A).

b. Radiograph envelope containing seeds.

c. Classify seed images on radiograph (see section

VI). (Record: columns 46-61)

Step 158

a. Take developed seeds from removed cone scales

and place in separate envelope.

b. Radiograph envelope containing seeds.

c. Classify seed images on radiograph (see section

VI). (Record: columns 62-77)

Step 16

Count any remaining second-year aborted ovules and add to number previously extracted (step 10). (Record: columns 41-42)

1 J

4

I

Step 17

Determine the number of first-year aborted ovules by subtracting

the total number of second-year aborted ovules plus the total number of developed seeds from two times the number of fertile scales

on the cone. (Record: columns 38-40)

Step 18

Place all developed seeds in germination test using standard germination testing procedures.

+

Step 19

Summarize data and compute important seed production indicator

values (see section VII). I

ZFirst-year aborted ovules may be counted as the scales are removed by observrng the location or the scale where the seed is produced .%afes wit

first-year aborted ovules do not have the depress/on formed in the scale where the seed was located

12
Figure 7.-(A) Shortleaf pine cone at harvest. (8) Same cone with lower infertile scales removed. Remaining scales are all fertile except

for a few infertile scales at the cone tip.

Figure 8.-Cone scale rerrIoval.

removal is: alternated withdrilli!

short dista‘rices of the,cone axis.
VI. CAUSES OF OVULE

AND SEED LOSSES

After first-year aborted ovules and second-year aborted ovules have been separated from developed seeds, the causes

for seed losses can be rdentrfred. The cause for abortron of an bvule often must be Inferred from the time at which abortion

occurred. Developed seeds are classrfred by the appearance of the embryo and gametophyte tissue on a radrograph (fig. 9).

Classes of ovules and seeds are defined. and causes for losses are summarized In tabular form In the Appendix.

FIRST-YEAR ABORTED OVULES

There are two causes of ovule abortron In the first Year ~ lack of pollen and seedbug damage

Lack of viable pollen. - Pollen IS an obvrous requirement for the production of healthy cones and viable seeds (Sarvas

1962) For productron of vrable seeds, two pollrnatron requirements must be met. Pollen must be available when female

flowers are receptive, and It must germinate and grow into the ovule tissue Female flower receptrvrty begins when the

opening between the scales IS large enough for pollen to enter and extends from a few days to more than a week. Once the

scales have closed, addrtronal pollen cannot enter the ovule. If no pollen IS present or If the pollen falls to germinate, the

ovule aborts early rn the first Year of cone development (Bramlett and Johnson 1975; McWrllram 1959, Sarvas 1962).

Seedbug feeding. ~ Feeding by nymphs of the leaffooted pine seedbug, Leptog@suscorculus (Say), often causes abortron of a high percentage of ovules (DeBarr and Kormanrk 1975; DeBarr and Ebel 1.973; DeBarr and others 1975) Ovules

destroyed by seedbugs cannot easily be drstrngurshed from ovules aborting from lack of pollen.

Losses to seedbugs can be estimated by protecting some cones in screened cages and comparing yields of protected

with unprotected cones Wrth thus technique, the impact of seedbugs can be separated from losses to physiologrcal and envirorimental factors

Figuire 9.-Classification of radiographic images of developed (A) filled (potentially sound] seeds; (8) mechanically

seeds; (C & DJ malformed seeds-C. incomplete gametophyte: D, distorted embryo (IeftJ; missing embryo (right); (continued)
Figure 9 (continued).-(EJ damaged by seedbug; (F) destroyed by seedworm; lGI destroyed by seed chalcid; (HI empty seeds and (I& JI

damaged by fungi

15
SECOND-YEAR ABORTED OVULES

Seedbug feeding. - In extensive studies of shortleaf (P. echinata Mill.) and loblolly (P. taeda L.) pines (DeBarr 1974a),

slash pine (P. elliottii Engelm. var. elliottii) (DeBarr and others 1975). and Virginia pine (P. virginiana Mill.) (Bramlett and

Moyer 1973). second-year ovule abortion has clearly been shown to result almost entirely from feeding by the leaffooted

pine seedbug in early summer (fig. 10). These ovules appear resinous, collapsed, or necrotic on the mature cone scales

(figs. 48; 5C. left; and 5D. right).

Developmental problems.-Some ovules stop developing during the second year after they have developed a seedcoat.

These “miniature seeds” may be oniy slightly larger than first-year aborted ovules or up to one-half the normal size for

seeds of the species. These are all empty. The cause of this type of ovule mortality is unknown, but It apparently results

from a block in the normal developmental sequence prior to fertilrzation. The frequency of second-year aborted ovules due

to developmental problems has been quite low in production seed orchards.

DEVELOPED SEEDS _’

Filled, partially filled, and empty developed seeds are readily distinguishable on radiographs. In addrtron, several types oi

damage can be identified on radiographs (fig. 9). :.

Figure IO.-Diagram of a cone showing how

the leaffooted pine seedbug pen&rates alternate layers of cone scales and seed wings to

puncture seeds lying along the cone axis.

(From DeBarr 1970)

Filled Seeds

Filled seeds (potentially sound seeds) are those that have healthy undamaged gametophyte tissue. a normal embryo.

and no evidence of insect or fungal damage. These seed structures are readily distinguished on a radiograph (fig. 9A).

Potentially sound seeds may be mechanically damaged at any stage of extraction (fig. 9B). Since the primary purpose of

cone analysis is to judge production efficiency in the seed orchards, mechanically damaged seeds should be classed as

filled.

Partially Filled Seeds

Seedbug.-The leaffooted pine seedbug and the shieldbacked pine seedbug, Tetyra bipunctata (H.-S.), feed on secondyear cones, damaging and destroying seeds (DeBarr 1967). Damage to fully developed seeds can often be identified on

radiographs by the appearance of the gametophyte tissue (DeBarr 1970). Typical seedbug feeding removes irregularly

shaped portions of gametophyte tissue (fig. 9E). Extended feeding on a single seed often removes most of the gametophyte

tissue and embryo, leaving only a shriveled remnant in the seedcoat.

16
Seedworm.-Three species of seedworms, Laspeyresia spp.. Infest seeds of the southern prnes (Ebel and others 1975).

Srnce seedworm-damaged seeds generally are stuck to the cone scales wrth excrement and sulk webbing, most are detected and counted as the rndividual cone scales are dissected during cone analysrs. The few seeds included In the extracted

sample are easily detected on the radiograph (fig. 9F). Seedworm larvae are not present in the seeds, and the exit holes are

large. The gametophyte tissue has been replaced by larvel excrement and bonng frass that looks granular on the radiograph

and often is formed Into concentric rings (DeBarr 1970)

Seed chalcid-A seed infested by a seed chalcrd, Megastigmus atedius Walker, lacks gametophyte tissue and contains a

srngle larva that is readily visible on the radiograph (fig. 9G). To date, the only prne growing In the Southern Unrted States

known to be infested by this seed chalcid is eastern whrte pine (P. strobus L.) (Speers 1975)

Fungi-Fungal damage IS recognizable on radiographs as (1) cloudy or hazy semrcrrcular areas generally at the margin of

the gametophyte trssue (fig. 91). (2) partial to complete disorganization of both the gametophyte and embryonic trssues (fig.

9J). and (3) all gametophyte tissue and embryo shrunken or “mummrfied.” Fungal da_mage may have the same appearance

as advanced seedbug damage on the radiograph. Dissectron and close examrnatron of rnfected seeds have revealed fungi In

the gametophyte trssues, embryo, or both. The mechanisms of fungal penetratron are cot known, but the presence of fungi

In seeds is known to reduce the germrnatron percentage and to increase damping-off of seedlings (Miller 1976).

Ma/formed seeds.Seeds with incomplete embryos, distorted embryos, or incomplete gametophytes are classed as

malformed. Missing or malformed embryos show up clearly on radiographs (fig. 9C and D). These malformatrons have

physiological causes. In some malformed seeds, the gametophyte trsse does not completely fill the seedcoat and no rnternal insect, disease, or mechanical damage is evident (fig. 9C). In these seeds, approxrmately one-half to three-quarters of

the normal amount of gametophyte tissue is normally present. Seeds with incomplete gametophytes occur in cones harvested before.maturity. Malformed seeds occur at a low frequency in most lots and rarely exceed 1 to 2 percent for most

trees.

Empty Seeds

Seeds that contain only a remnant of gametophyte or embryo tissue (fig. 9H) are classed as empty. In the trade. empty

southern pine seeds are often called “pops.” There are two known causes of empty seeds. First, each pine parent carries

some recessive lethal genes (Bramlett and Pepper 1974; Franklin 1969). When two recessive lethal genes for the same

trait are present, the embryo dies and the gametophyte tissue does not develop. The result IS an empty seedcoat with only

a remnant of the dead embryo present. Second, seedbug feeding during the early stages of embryo and gametophyte

development will produce a completely empty seedcoat. The gametophyte tissue is completely destroyed by digestive enzvmes of the insect.

VII. SUMMARIZING DATA

To interpret the information obtained from the cone analysis, the data must be summarized. The summary should be

tabulated according to trees, clones, or other categories of interest, For the following items, a numencal average can be

computed:

Cone opening (CO)

Length (LT)

Lower infertile scales (LI)

Filled seeds (FL)

Malformed seeds (MS)

Seedbug-damaged seeds (SB)

17
Upper infertile scales (UI) Seedworm-destroyed seeds (SW)

Fertile scales (FS) Seed chalcid-destroyed seeds (SC)

First-year aborted ovules (Al) Fungal-damaged seeds (FN)

Second-year aborted ovules (A21 Empty seeds (EM)

Extracted seeds (ES) Germinating number of seeds (GN)

From the basic summary of the raw data, the key indicators of seed performance are calculated as shown below. Numbers In parentheses are the columns on the suggested data form in the Appendrx.

1 .

2.

3.

Indicator _’ Computation

Seed potential (SP) = 2 x Fertile scales or 2 x (3436)

Total developed seeds (TS) = Seedbug-damaged seeds + seedworm-destroyed seeds + seed

chalcid-destroyed seeds + fungal-damaged seeds + malformed seeds + empty seeds + filled seeds or

(4647 and 62-63) + (48-49 and 64-65) + (50-51 and 66-67) + (52-53 and 68-69) + (54-55 and 70-71)

+ (56-58 and 72-74) + (59-61 and 75-77)

Percent developed seeds = Total developed seeds x 1oo .,T$ x 1oo

Seed potential

4. Percent filled seeds = Filled seeds

Total developed seeds

x 1oo or (59-61 and 75-77) x TOO

TS

5. Percent known insect-damaged seeds =

Seedbugdamaged seeds + seedworm-destroyed seeds +

seed chalcid-destroyed seeds x 100 o r

Total developed seeds

(4647 and 62-63) + (48-49 and 64-65) + (50-51 and 66-67) x TOO

TS

6. Percent malformed seeds = Malformed seeds

Total developed seeds

x lOOor{ 54-55 a;: 70-71) x 1OO

7. Percent fungal-damaged seeds = Fungal-damaged seeds x 1oo or (52-53 and 68-69) x 1OO

Total developed seeds TS

8. Percent first-year aborted ovules = First-year aborted ovules x 1oo or (38-40) x TOO

Seed potential SP

9. Percent second-year aborted ovules = Second-year aborted ovules x TOO or (41-42) x 100

Seed potential SP

10. Seed efficiency (SE) = Total filled seeds or (59-61 and 7577) x 1OO

Seed potential SP

18
11, Extraction efficiency (EE) =

Procedure A: Extracted developed seeds

Total developed seeds

orvx 100

Procedure B: Extracted filled seeds (59-61)

Total filled seeds 0r(59-61 and 75-77) ’ loo

12. Germination percentage (GP) =

Procedure A: Germinated seeds x 1oo or (78-80)

Total filled seeds (59-61 and 75-77) ’loo

Procedure B: Germinated seeds x 100or~x 100 -.

Extracted filled seeds ,’

13. Seedling efficiency = Seed efficiency x Extraction efficiency x Germinatidn percentage or SE x EE x GP

10,000 10,000

VIII. -DATA INTERPRETATION

AND UTILITY

The purpose of cone analysis is to evaluate seed production and to identify when and why potential seeds are lost. Table

1 shows typical values for key indicators in a well-managed seed orchard. It also shows values that may indicate need for

corrective action.

The seed potential defines the biological limit for the number of seeds produced by each cone. Thus, each tree species

has an average seed potential and a range of observed values based upon the number of fertile scales per cone. For example, the seed potential for cones from a slash pine seed orchard averaged 170. Loblolly pines had an average seed potential

of 155, and Virginia and shortleaf pines had averages of 88 and 87 (Bramlett 1974). Variation In seed potential IS relatively

small within specific clones. Seed potentials in a given orchard, however, may differ from the average for the species,

depending on the particular clones in the sample.

Aborted ovules are potentral seeds that die before the formatron of a normal seedcoat. For all the southern pines, ovule

abortion can be a serious cause of low seed yields The two known causes of first-year aborted ovules (lack of pollen and

seedbug damage) are influenced by the orchard age. In young orchards, female flowenng of many clones usually begrns

before male catkin productron is adequate to ensure complete pollrnatron. Unless there IS a supply of pollen from surrounding stands, many ovules die within 2 months from lack of viable pollen. In contrast, older seed orchards have an

abundant pollen supply but also often have large seedbug populations that cause extensive frrst-year ovule abortion in unprotected conelets. Conclusive separation of Insect from pollination problems requires comparison of full seed yields from

cones with and without screened cages that exclude Insects.

Second-year aborted ovules are, for practical purposes, a direct indication of seedbug feeding. Other agents seldom

cause second-year abortrons exceeding 1 to 2 percent of the seed potential. Second-year ovule abortion may reach 50 percent, but seedbug control may be warranted when the loss approaches 10 percent of the seed potential To prevent

second-year ovule abortion, insectrcrde applications must be timed to reduce populatrons of leaffooted pine seedbugs dur19
rng the sprrng when the conelets are rapidly enlargrng to full-srzed cones. Seedbug feedrng on full-sized cones causes

empty or damaged seeds rather than aborted ovules

Total developed seed IS srmply the number of normal, full-srzed seeds In each cone. Obvrously. the orchard manager

strives to maxrmrze developed seed productron Wrth adequate pollrnatron and effectrve insect control. the developed seed

yields should approach 80 to 90 percent of the seed potential

Yields of developed seeds are meanrngless wrthout considering seed quality. Using radiography. the actual number of

frlled seeds per cone IS determrned Then seed efficiency IS expressed as the ratro of filled to potentral seed. Thus. seed effrcrency measures the productrvrty of a cone In relation to Its brologrcal capacrty The average seed efficiency for a clone or

an orchard IS the single most Important measure of seed production It can be used to compare the relative performance of

two pine specres wrth drfferent seed potentrals For example, If a shortleaf pine produces an average of 45 frlled seeds out

of a potentral of 90, the seed efflclency would be 50 percent However, 45 filled seeds In a slash pine cone with a Ventral

of 180 lndlcates a seed efflclency of only 25 percent

TABLE 1. - KEY INDICATORS FOR DATA INTERPRETATIGN

Cone characteristic and

threshold level (percent) 1 Interpretation

1. First-year aborted ovules

O-10 Good management practice

11-19 Evaluate possible insect or pollination problems

20+ Identify and correct insect or pollination problems

2. Second-year aborted ovules

o-5 - Good management practice

6-9 Evaluate seedbug control program

10+ Increase seedbug control

3. Percent filled seeds

B5+ Good management practice

50-84 Evaluate possible seedbug or inbreeding problems

Below 50 Identify and correct seedbug or inbreeding problems

4. Insect-damaged seeds

O-10 Good management practice

11-19 Evaluate insect problem and control program

20+ Improve or increase control program

5. Seed efficiency

55+ Good management practice

35-54 Evaluate causes of seed loss

Below 35 Identify causes of seed loss and begin correction

6. Extraction efficiency

90+ Good management practice

Below 90 Identify and correct improper conditions or immature cone collections

7. Germination percentage

90+ Good management practice

80-89 Identify cause (May be: undetected seedbug damage or mishandling of seed)

Below 80 Identify and correct problem (May be: improper germination, undetected

seedbug damage, mishandling of seed)

8. Seedling efficiency

50+ Good management practice

26-49 Identify cause - poor germination, immature cones

Below 25 Identify and correct cause

’ Each organrzatron may wash to modify these thresholds to meet Its targets

2 0
The brologrcal maximum for seed effrcrency may be as hrgh as 80 percent for rndrvrdual cones, but values thus high are

seldom observed under seed-orchard conditrons. Seed efficrencres greater than 70 percent have occurred when screenwire cages were used to protect slash pine cones from Insects (DeBarr and others 1975)

Seed efficiency values of 35 to 54 percent rndrcate that some potentral seeds are being lost. Yrelds could be Increased

by rdentrfyrng the specific causes of seed losses and prescnbrng corrective procedures wrthrn economrc constraints Seed

effrcrencres of less than 35 percent rndrcate excessrve losses The causes should be Identified and the problems corrected

as soon as possrble

Developed seeds not classified as “frlled” are worthless Thus, filled seed percentage srmply expresses the ratro of filled

to developed seeds Filled seed percentages seldom exceed 90 percent In commercral operatrons, but greater values have

been reported wrth controlled pollrnatrons or effective seedbug control (Bramlett and Pepper 1974, Merkel and others

1976) Wrnd-pollrnated cones from orchards should average 85 percent or more filled seeds Lower values Indicate losses

In some or all of the classes rdentrfred from the radiographs Indrvidual-loss classes exceedrng 10 percent warrant corrective

measures _.

Insects. particularly seedbugs, destroy a large percentage of the developed seeds In,cone analysrs, losses are attributed

to seedbugs, seedworms, and seed chalcrds. Usually, some seeds classed as “empty” are also the result of seedbug

feedrng Therefore, as recognized seedbug losses Increase. the number of empty seeds often rises (DeBarr 1974b. DeBarr

and Ebel 1973).

Malformed seed losses (Incomplete gametophyte tissue, distorted embryo, missing embryo1 seldom exceed 10 percent

of the developed seeds In well-managed orchards. Incomplete gametophytes apparently result from retarded development

or Inadequate nutrition. Cones collected before maturity produce seeds with incomplete gametophytes. Malformed seeds

occur In sample lots at such low frequencies that these types of losses are of only minor importance.

Fungus damage to southern pine seeds has only recently been recognized as a potentral problem. The Importance of

fungal damage IS undetermined. Part or all of the seeds from cones with external symptoms of fungal damage may be infected by fungi. Correctrve measures are not yet available to prevent seed destruction by fungi.

The causes of empty seeds are difficult to identify Lethal genes In the embryo, seedbugs. and possrbly fungi cause

empty seeds. In wind-pollrnated seeds from an orchard, as many as 15 percent of the developed seeds may be empty This

value would reflect some effects of self-pollrnatron wrthrn the clones In the orchard (Kraus 1975); the proportron of empty

seeds IS normally high after self-pollinatron. Lethal genes In embryos cannot be elrmrnated easrly Thus genetrcally controlled

system mrnrmrzes Inbreeding and favors outcrossing In pines

If more than 15 percent of the developed seeds are empty, seedbug damage should be suspected and control programs

should be reevaluated. Seedbug damage can be reduced with Insecticides.

Extraction efficiency measures the ease with which developed seeds are removed from the cones. It IS the percentage of

developed seeds extracted by kiln or air-drying the cones. It should be noted that this extraction IS done In a laboratory -

that the results are not necessarily the same as would be obtained by commerctal extraction. Percentages below 90 Indicate poor extraction technique or immature cones.

Germination percentage measures the vrabrlity of seeds classrfred as “filled” (potentrally sound). This value normally

averages 90 percent or better Lower germrnatron percentages may Indicate Improper classrfrcatron of some seeds In the

lot (Rowan and DeBarr 1974). Germination percentages are useful for comparing the vrabrlrty of collectrons from different

trees or from subsequent years.

Seedling efficiency is simply the product of the three efficiency values - seed efficiency, extraction efficiency, and germination percentage. It reflects the total yields of the cone, seed, and seedling crops from the orchard. If any one of the efficiency values IS low, the overall effrcrency of the orchard IS reduced. A maximum realistic goal for production orchards IS a

seedling efficiency of about 55 percent. Such would be the case if seed efficiency = 70 percent, extraction efficiency = 90

percent, and germrnatron percentage = 90 percent.

Cone analysrs does not estimate all seed losses; it only measures seed losses from surviving cones. Cone losses from

the time of flower Initiation to cone maturity are not considered. Often 50 percent or more of the seed crop IS lost during

this period. By combrnrng cone crop life tables with cone analysis, the total seed production for an orchard can be evaluated

(Fatzinger 1975; DeBarr and Barber 1975; Bramlett 1972a; Ebel and Yates 1974).

21
lx. FOR ADDITIONAL INFORMATION

For help with known insect problems or assistance in identifying insect problems, contact:

STATE FOREST ENTOMOLOGIST

Alabama Forestry Commission Louisiana Forestry Commission

5 13 Madison Ave. P.O. Box 1628

Montgomery, AL 36104 Baton Rouge, LA 70721

205/832-5897 5041389-7 12 1

Tennessee Division of Forestry

2611 West End Ave.

Nashville, TN 37203

615/741-3326

Arkansas Forestn/ Commission

3821 W. Roosevelt Rd.

Little Rock, AR 72204

501/371-1736

Mississippi Forestry Commission

908 Robert E. Lee Building

Jackson, MS 39201

601/354-7124

Florida Division of Forestry

Collins Building

Tallahassee, FL 32304

904/488-7936

Georgia Forestry Commission

P.O. Box 819

Macon, GA 3 1202

9 121744-3241

Kentucky Division of Forestry

207 Holmes St.

Frankfort, KY 40601

502/564-4496

N.C. Dep. of Natural Resources

P.O. Box 27687

Raleigh, NC 2761 1

919/733-4141

Oklahoma Forestry Division

Capitol Building

Oklahoma City, OK 73 105

405152 l-3886

S.C. State Commission of Forestry

P.O. Box 2 1707

Columbia, SC 29202

803/758-226 1

Texas Forest Service

P.O. Box 310

_. Lufkin, TX 75901

7 131632-7761

Virginia Division of Forestry

P.O. Box 3758

Charlottesville, VA 22903

8041977-6555

FOREST INSECT AND DISEASE MANAGEMENT ZONE OFFICE

Forest Insect & Disease Management

Seed Orchard and Nursery Insects

P.O. Box 5895

Asheville, NC 28803

7041258-2850 x625

Forest Insect & Disease Management

Seed Orchard and Nursery Insects

2500 Shreveport Highway

Pineville, LA 7 1360

318/445-6511

For questions on the use of pesticides, consult:

Pesticide Specialist

Resource Protection

Suite 706

1720 Peachtree Rd.,

Atlanta, GA 30309

404188 l-3734

NW

For further information on cone analysis or on the service supplied, contact:

Eastern Tree Seed Laboratory

P.O. Box 819

Macon, GA 3 1202

9 12/744-33 11

22
X. GLOSSARY

Many of the deflnltlons given below are based on those In the ‘Glossary for Forest Tree Improvement Workers’ (Snyder 1972) and

‘Seeds of Woody Plants In the Unlted States” (USDA Forest Service 19741

ABORTED OVULE:

FIRST-YEAR Any potential seed that aborts (ceases development) during the first growing season or conelet stage of pine seed

productlon

SECOND-YEAR Any potential seed that aborts (ceases development) during the second growing season before fertlllzatlon or seedcoat hardening

ARCHEGONIUM: Female sex organ In the ovule of pine

CATKIN: The male strobllus which produces pollen

CLONE: Group of genetlcally Identical plants produced by vegetatively propagating a single plant over one or more generatlons. accomplished by grafting, rooting, air-layering. or tissue culture Compare ortet. ramet _’

CONE: The female reproductive structure of pines. conslstlng of a central axis supportlng bract?. each of which subtends a scale bearIng naked seeds :.

CONE ANALYSIS: A systematic procedure to evaluate the seed productlon potential and efflcleney of pine cones

CONE AXIS: Central core of the cone

CONELET: Immature female cone of pines from time of female “flower” scale closure after polllnatlon until the lnltlatlon of rapid

development of the cone a few months b-fore maturity

COTYLEDONS: First leaves developed In the embryo of a seed They become functlonal leaves after germlnatjon. Immature leaves

FEMALE CONE: See cone

FEMALE FLOWER: The pine female strobllus (cone) pnor to polllnatlon

FERTILE SCALE: A cone scale that IS capable of producing seed, generally Includes most scales In the upper one-half to two-thirds of

the cone

FERTILIZATION: The union of the egg (1 N) wlthln the ovule and the sperm (1 N) from the pollen tube to form the embryo 12N)

FILLED SEED (POTENTIALLY SOUND) Mature ovule contalnlng an embryo and nutntlve tissue enclosed In layers of protective tissue

(seedcoat), capable under suitable condltlons of development Into a plant slmllar to the one that produced It

GAMETOPHYTE TISSUE: Haplold material (1 N) surrounding the embryo, provides nutntlonal reserves for the seed of pine

GERMINATION: Sufflclent development of plant parts from a seed to indicate a potential for developing Into a normal plant

HYPOCOTYL: All of the axis of an embryo or stem of a seedling between the cotyledons and the radlcle. an immature stem

INFERTILE SCALE: A pine cone scale Incapable of producing seed, located at the base and apex of the cone

INTEGUMENT: A single layer of tissue encloslng the pine ovule and nucellus After fertlllzatlon. It develops Into the seedcoat

MICROPYLE: Small opening In the Integument of an ovule through which the pollen grain normally passes to reach the pollen chamber

NUCELLUS: Mass of thin-walled cells that composes the central and maln part of the body of an ovule. It IS surrounded by the Integument

OVULE: A female structure which contains an egg cell and develops Into a seed

POLLEN GRAIN: The winged male fertlllzlng structure

POLLEN TUBE: An elongate outgrowth of the germlnatlng pollen grain.

POLLINATION: The transfer of pollen to the receptive female flower

POTENTIALLY SOUND SEED: Seed fllled with apparently undamaged tissue on which vlablllty has not been tested

RADICLE: Portion of the axls of an embryo from which the root develops. Immature root

RAMET: Independent member of a clone

SEED: “In conifers” the structure formed after fertllzatlon: lnltlally consists of seedcoat, embryo, and gametophyte storage ttssue

SEEDCOAT: Protective layer on a seed derived from an Integument

SEED ORCHARD: A plantation conslsttng of clones or seedlings from selected trees and cultured for early and abundant seed

productlon

SEED POTENTIAL: Two times the number of fertile cone scales or the maximum number of seeds a cone IS capable of producing

STROBILUS: Male or female frultlng body of pines

23
Xl. LITERATURE CITED

Bramlett. D L

1972a Cone crop development records for SIX years in shortleaf pine For SCI 18 31-33

Bramlett D L

1972b Seed and aborted ovules from cones of P/nus echmara Ga For Res Count Res Pap 7 1 4 p

Bramlett. D L

1974 Seed potential and seed efflclency. p l-7 ln John Kraus (ed ), Seed yield from southern pine seed orchards colloquium proceedings Ga For

Res Count, Macon. Ga

Bramlett. D L and C J Johnson

1975 Polllnatlon or- a slash pine seed orchard Thirteenth South For Tree lmprov Conf Proc 1975 161~ 165

Bramlett. Cl L. and E L Moyer Jr

1973 Seed losses reduced in Vlrglnla pine cones by screen wire cages USDA For Serv Res Note SE-1 92 4 p Southeast For Exp Stn,

Asheville. N C

Bramlett. D L. and W D Pepper

1974 Seed yield from a dlallel cross or- Vlrglrxa pine. p 49-55 In John Kraus ied j Seed yleld frqm southern pine seed orchards colloquium

proceedings Ga For Res Count, Macon, Ga

,’ DeBarr. G L

1967 Two new sucking insect pests of seed in southern pine seed orchards USDA For Serv Res r\jbte SE-78. 3 p Southeast For Exp Stn

Asheville. N C

DeBarr. G L

1970 Charactenstlcs and radIographIc detectlon of seedbug damage to slash pine seed Fla Entomol 53 109- 1 17

DeBarr. G L

1974a Conelet abortIon. ovule abortjon. and seed damage by Leptoglossus corculus (Say) in squthern pine seed orchards Ph D DISS Dep Entomol Unlv Ga Athens 189 p

DeBarr. G L

1974b Quantlfyng the Impact of seedbugs. p 34-41 In John Kraus (ed 1, Seed yield from southern pine seed orchards colloquium proceedings

Ga For Res C&nc, Macon Ga

DeBarr. G L, and L R Barber

1975 Mortality factors reducing the 1967~ 1969 slash pine seed crop in Baker County. Florlda a life table approach USDA For Setv Res Pap

SE-131. 16 p Southeast For Exp Stn Asheville. N C

DeBarr G L. D L Bramlett. and A E Squlllace

1975 Impact of seed (“sects on control-pollinated slash pane cones Thirteenth South For Tree lmprov Conf Proc 1975 177-181

DeBarr. G L. and B H Ebel

1973 How seedbugs reduce the quantity and quality of pine seed ylelds Twelfth South For Tree lmprov Conf Proc 1973 97-103

DeBarr. G L. and P P Kormanlk

1975 Anatomical basis for conelet abortjon on Plnus echmata followlng feedlng by Leptoglossus corculus (Hemlptera Coreldae) Can Entomol

107 81-86

Ebel. B H T H Flavell L E Drake H 0 Yates Ill, and G L DeBarr

1975 Seed and cone Insects of southern pines USDA For Serv Gen Tech Rep SE-8, 40 p Southeast For Exp Stn Asheville. N C, and

Southeast Area, State and Prlv For, Atlanta. Ga

Ebel. B H, and H 0 Yates Ill

1974 Insect-caused damage and mortality to conelets. cones, and seed of shortleaf pine J Econ Entomol 67 222-226

FatrInger. C W

1975 Seed orchard sampling commlttee Minutes 20th South For Insect Work Conf Baton Rouge. La p 41-43

FrankIln E C

1969 InbreedIng depression or- metrlcal traits of loblolly pine iP/nus taeda L i as a result of self-polllnatlon N C State Unlv Sch For Resour Tech

Rep 40. 19 p

Karrfalt. R P. and E W Belcher. Jr

1977 Evaluation of seed production by cone analysis 24th Northeast For Tree lmprov Conf Proc 1976 84-89

Kraus. J F

1975 Estimates of selfed seedling productlon from a slash pine seed orchard based on gene markers Thirteenth South For Tree lmprov Conf

Proc 1975 93-96

Lyons L A

1956 The seed production capacity and efflclency of red pine cones iF’!nus resinosai Can J Bot 34 27~36

McWtlllam J R

1959 InterspecIfIc lncompatlblltty in Pmus Am J Bot 46 425~433

Merkel E P G L DeBarr and C H O’Gwynn

1976 Mist blower appllcatlon of Guthlon” for cone Insect control 117 slash pine seed orchards USDA For Serv Res Pap SE 148 8 11 Southeast

For Exu Stn Asheville N C

24
Miller. T

1976 The assoclatlon of fungi with cone and seed losses in the southern pines (Abstr J In Proceedings forest tree seed testing and handling

workshop. Macon. Ga USDA For Serv, Southeast Area, State and Pnv For 1 p

Rowan. S J and G L DeBarr

1974 Moldy seed and poor germlnatlon linked to seedbug damage In slash pine USDA For Serv Tree Plant Notes 2511 i 25-27

Sarvas. R

1962 lnvestlgatlons on the flowering and seed crop of P/nus s/lvestr/s. Commun lnst For Fenn 53 198 p

Snyder. E B

1972 Glossary for forest tree Improvement workers USDA For Serv, South For Exp Stn 22 p New Orleans. La

Speers. C F

1975 Megastlgmus atedm a newly dtscovered pest of eastern white pine seed In North America Ann Entomol Sot Am 68 559-560

U S Department of Agriculture. Forest Service

1974 Seeds of woody plants In the Unlted States U S Dep Agnc Agnc Handb 450 883 p

25
XII. APPENDIX

ITEM

Test number

Year

Species

Orchard

Clone

Ramet

Cone number

Cone opening

Cone length

CONE ANALYSIS DATA CARD CODING AND COLUMN NUMBERS

Lower infertile scales

Upper infertile scales

Fertile scales

First-year aborted ovules

Second-year aborted ovules

Extracted seeds

Soedbug-damaged seeds

Seedworm-destroyed seeds

Seed chalcid-destroyed seeds

Fungal-damaged seeds

Malformed seeds

Empty seeds

Filled seeds

Germinated seeds

CODE COLUMN

TN

Y R

SP

OR

CL

Identification

Input

1q-17

RA

CN

c o 26 1.

LT 29-31

LI 32-33

UI 37

Optional

Data

FS 34-36

A l 36-40

A 2 41-42

ES

SB

SW

SC

FN

M S

EM

FL

GN

26

43-45

46-47.62-63

48-49.64-65

50-51,66-67

52-53.68-69

Developed

Seeds
iQ

27
CLASSIFICATION OF OVULES AND SEEDS

Columns on data card

Class Procedure A Procedure B Identity1 Cause Comment

A. Aborted ovules

1. First-Year 38-40 3840

2. Second-Year 41-42 41-42

B. Developed seeds

1. Filled

2. Partially filled

Seedbugdamaged

E

Fungaldamaged

Malformed 54-66

3. Empty

Seedwormdestroyed

Seed chalciddestroyed

Empty 56-58

59-61

46-47

52-53

48-49

50-51

Small undeveloped ovules with a. Lack of viable pollen Requires caged cones to

normally developedlwing (V) b. Seedbug feeding establish cause

Flattened seed often a. Seedbug feeding Usually due to seedbug

resinous (V) b. Developmental problems attack

69-61

75-77

Full-sized seed with complete includes mechanically

seedcoat (VI, embryo, and a damaged filled seed;

gametophyte that fills the does not change

seedcoat (RI classification

46-47

62-63

52-63

68-69

Irregular areas of missing

gametophyte tissue (R)

Seedbug feeding

Cloudy or hazy semicircular areas

at margin of gametophyte tissue,

disorganization or shrunken or

mummified gametophyte tissue (RI

Fungi --

64-65

70-71

Missing or damaged embryo or

gametophyte tissue does not fill

seedcoat (RI

Physiological Early cone harvest will

cause incomplete gametophyte tissue

48-49 Seed filled with granular

64-66 material (R) Seedworm feeding

Entry-exit hole in seedcoat (V)

50-51

66-67

Larva in seedcoat (R) Seed chalcid feeding Limited to eastern white

pine seed

66-58 Seedcoat empty or containing a a. Seedbug feeding Requires caged cones to

72-74 shriveled remnant of tissue (RI b. Recessive lethal genes establish cause

lThe deslgnatlon “V” mdlcates ldentlfymg character vwble to unaided eye. “R” Indicates vwble only on radiograph.

I

Southeastern Area S. 6 P. F - 8