Scientific Reports of the
Hokkaido Salmon Hatchery
No. 49: 1-9, March 1995

Variation in Body Size, Fecundity, and Egg Size of
Sockeye and Kokanee Salmon, Oncorhynchus nerka,
Released from Hatchery
Masahide Kaeriyama, Shigehiko Urawa and Masa-aki Fukuwaka
Research Division, Hokkaido Salmon Hatchery, Fisheries Agency of Japan,
2-2 Nakanoshima, Toyohira-ku. Sapporo 062, Japan

Abstract. - Adult sockeye salmon (Oncorhynchus nerka) returning to the Bibi River, which were originally derived from LakeShikotsu kokanee salmon,and adult kokanee salmon in Lake Shikotsuwere examined for variation in body size, fecundity, and egg size. Both groups of adults originated from hatcheryreleasedjuveniles. Larger adult females had higher fecundity. Fecundity and fork length fitted allometricformula within each population. There was no relationship between fork length and egg size in Lake
Shikotsu kokanee salmon or in age-I . 1 sockeye salmon from the Bibi River although the sockeye salmon
were approximately 65% larger in fork length than the kokanee salmon. Age-1.2 sockeye salmon, however, had eggs about 12% larger than did kokanee and age-1.1 sockeye salmon. We suggest fecundity inhatchery-releasedsockeye salmon vary with body size influenced by environmentalcomponents, and that
their egg size would be affected by both environmental and genetic components such as polygene model
within a cohort or population because of exclusion from the breeding competition and parental care.

body size contributed to adult female fitness such as
biomass of egg production (Holtby and Healey, 1986;
van den Berghe and Gross, 1989; Beacham andMurray, 1993), and a significant positive relationshipbetween body size and egg production (fecundity andegg size) has been observed among Pacific salmon
populations (Watanabe, 1955; Beacham and Murray,
1986, 1988; Beacham et al., 1988; Fleming and Gross,
1990; T a l h a n and Healey, 1991).
Relative distance of freshwater migration to
spawning grounds generally has a marked effect on
both fecundity and egg size, and populations spawningin the upper portions in drainages of large rivers have
reduced fecundity and egg size compared with coastal
spawning populations (Beacham and Murray, 1993).
Generally, fecundity is higher, and egg size is lower inmore northern populations of Pacific salmon (Flemingand Gross, 1990; Beacham and Murray, 1993).
Timing of spawning in Pacific salmon can have agenetic component (Bams, 1976; Taylor, 1980).
Within an area, early-spawning populations have high-Introduction
An anadromous form of sockeye salmon(Oncorhynchus nerka) is not naturally found in Japan,
although the lacustrine form (kokanee salmon)
released from hatcheries are observed in several oligotrophic lakes. Lake Shikotsu kokanee salmon were
derived from kokanee eggs from Lake Akan on
Hokkaido in 1893 and sockeye eggs from Lake
Urumobetsu on Iturup Island during 1925-1940. The
population in Lake Shikotsu has been geographicallylandlocked in this lake for more than 15 generations.
Anadromous sockeye salmon have been produced
from Lake Shikotsu kokanee salmon by smolt releasetechnology in the Bibi River of the Abira River system, central Hokkaido (Kaeriyama, 1989, 1991,1992;
Urawa, 1991; Kaeriyama et al., 1992).
Among Pacific salmon, female sockeye salmon
have the highest fecundity and the smallest egg size
per given size of fish (Burgner, 1991). For sockeye
salmon, mean fecundities range from about 2,000 to
5,000 eggs female-'. Fecundity in kokanee salmon ismuch lower and may range from about 300 to lessthan 2,000 eggs female-' (Foerster, 1968; Burgner,
1991). Several studies have presented evidence thater proportions of older and larger individuals thanlate-spawning ones (Beacham and Murray, 1987).
The seasonal decrease in egg size coincides with aseasonal decrease in the body size of spawners, andthen progressively smaller fish lays progressively
smaller eggs in Pacific salmon populations (Beacham

Contribution A No. 346 from the Hokkaido Salmon Hatchery.

-1-a l l . bT-&&f$$i
SCI. REP.HOKKAIDO SALMON HATCHERY, NO. 49,1995

determine age. In addition, eggs were stripped fromeach female, weighed to calculate a gonad somatic
index (GSl), and fertilized by sperm. The following
equation was used to determine GSI: GSI =GSW/BW X 100, where GSW is gonad somaticweight (g), and BW is wet body weight. Eggs, isolated
from each female, were subsequently reared in Atkin's
incubator trays. At eyed egg stage, the total numberof eggs per female was counted, and egg size wasdetermined from volume (ml) of live water-hardened
eggs with excess surface moisture removed as follows:
D,= 20 (4Ve/(37rNe))"', where D , is egg diameter(rnm), V, and N, are volume and number of total alive
water-hardened eggs, respectively (Watanabe, 1955).
Relationships between body size and breeding
characters were analyzed using the following allometric regression (log-transformed linear regression): B, =aL b , where B , is fecundity (eggs female-') or egg
diameter (mm), L is fork length (mm), a is initial
growth constant, and b is a relative growth coefficient.

and Murray, 1987; Beacham et al., 1988).
We examine factors affecting fecundity and egg
size in Lake Shikotsu kokanee salmon and Bibi Riversockeye salmon.

Materials and Methods
Beginning with 1983-brood year eggs, Lake
Shikotsu kokanee salmon were reared at Chitose
Hatchery and then released as yearling smolts (age1.0) in the spring of 1985 in the Bibi River of the
Abira River system (Fig. 1). Anadromous sockeye
salmon have returned to the Bibi River as the first (F1 ) and second (F-2) filials at age 1.1 or 1.2 since 1986
(Kaeriyama, 1992; Kaeriyama et al., 1992).
To study variation in female body size, fecundity,
and egg size, we examined age-3.0 adult of kokanee
salmon in Lake Shikotsu during 1988-1994 and adultsockeye salmon in the Bibi River during 1990-1993(Fig. 1). Fork length (mm) and wet body weight (8)of females were measured and scales were collected to

N

f
River

-

0

/

5 km

Fig. 1. Locations where kokanee and sockeye salmon were sampled. B, the Bibi River of the Abira River system;
S, Lake Shikotsu of the Ishikm River system; S h , Shikotsu Hatchery where kokanee salmon eggs were reared;
C h , Chitose Hatchery where sockeye salmon eggs were reared.

2-

KAERIYAMA ET A L - REPRODUCTIVE CHARACTERS OF SOCKEYE AND KOKANEE SALMON

To analyze differences in body size, GSI, fecundity, and egg size among kokanee salmon, age-1.1 andage- 1.2 sockeye salmon populations, the Bartlet
method was first used to determine whether varianceswere equal. The analysis of variance (ANOVA) andthe nonparametric Kruskal-Wallis test were used tocompare populations. To determine differences in egg
size between populations, the least significant difference (LSD) method was also used.

in the 1988-1994 adult female kokanee salmon (Fig.
2A).
However, both GSI and egg size (egg diameter)
did not show annual changes, and remained stable atabout 17% and 5 mm respectively during 1988-1994(Table 1). There was no relationship between the fork
length and the egg size of Lake Shikotsu kokanee
salmon during 1988-1994 (n = 67, r2< 0.001, P > 0.5;
Fig. 2B).

Results

Variations in body size, fecundity, and egg size ofanadromous sockeye salmon
Table 2 shows fork length, GSI, fecundity and egg
size of Lake Shikotsu kokanee salmon in 1988-1994
and of Bibi River sockeye salmon in 1990-1993.
There are significant differences between kokanee
salmon, age- 1.1 and age- 1.2 sockeye salmon conceming the body size and fecundity (Kruskal-Wallis test,
P < 0.001). However, there was no difference between
any groups in GSI which remained at about 17%(Kruskal-Wallis test, P > 0.05). Significant difference(ANOVA,P< 0.001) in egg size was observed among
populations. Although there was not any difference ofegg size between kokanee salmon and age-1.1 sockeye
salmon (LSD test, P = 0.708), age-1.2 sockeye salmon

Annual changes in body size, fecundity, and egg sizeof kokanee salmon
Fork length of adult female kokanee salmon in
Lake Shikotsu averaged 215 mm during 1984-1988(Kaeriyama, 1991), showed increasing trend since
1989, and attained to 288 mm (SD = 8) in 1994 with
32% larger than in 1988 (Table 1). Fecundity also
increased with body size since 1989. Mean fecundity(666 eggs female-') of 1994 age-3.0 adult females
was twice than that (316 eggs female-') of 1988
females (Table 1). Fork length and fecundity weresignificantly related (n = 70, r2= 0.6463, P < 0.001); F= 0.00059 L24399,
where L and F are the fork length(mm) and the fecundity (eggs female - I ) respectively

Table 1. Annual changes in breeding characters of adult female kokanee salmon in Lake Shikotsu during 1988-1994. Average, standard error,
and number of individuals are presented by Ave, Se, and n, respectively.
1988

1989

1990

1991

1992

1993

1994

Ave
Se
n

219
0.8
156

244
0.4
940

264

252

248

249

288

0.8
200

0.6
250

0.7
190

0.5
90

0.5
141

Ave
Se
n

Ill

168
0.8
940

202
1.8
200

174
1.1
250

159

1.4
156

1.3
190

176
1
190

280
1.9
141

GSI (90)

Ave
Se
n

16.9
0.44
10

17.7
0.82
10

18.9
0.54
10

17.5
0.7
10

18.7
1.52
10

17.8
0.54
10

17.1
0.54
10

Fecundity(eggs female ')

Ave
Se

316
9.5

330
15.8

40 I
11.4

454
18.3

412
27.8

435
28.1

666
31

n

10

10

10

10

10

10

10

Ave
Se
n

4.8
0.06
10

5.2
0.06
10

5.1
0.03
10

5
0.03
10

5
0.03

5.1
0.09
10

4.8
006
10

Year
Fork length(mm)Body weight(d

Egg diameter
(mm)

-3-

10

dl? . ~ T W SCI
T ~REP HOKKAIDO SALMON HATCHERY, NO. 49,1995

I
-

e

*

8oo
700

.-> 600

-

-0

c

3

0

500 -

a>

LL

400 300 200

1
'

100
180

n = 70, 6 = 0.6463, P < 0.001
I

I

I

I

I

I

200

220

240

260

280

300

320

Fork length (L, mm)
6.0

BE- 5.5
E

m tt

. e

4.5

4.0

180

n = 67, P <0.001, P
I

I

200

220

I

I

240
260
Fork length (L, mm)

I

I

280

300

0.5

320

Fig. 2. Allometric relationships between fork length and fecundity (A), or egg diameter (B) of kokanee salmon in
Lake Shikotsu during the 1988-1994 spawning seasons.

had eggs approximately 12% larger than did kokanee
length (mm), fecundity (eggs female I), and egg
and age-1.1 sockeye salmon (LSD test, P < 0.001).
diameter (mm) respectively in the 1990-1993 adult
Relationships between fork length and fecundity
female sockeye salmon (Fig. 3).
(n = 37, r2 = 0.5520, P < 0.001), and between fork
length and egg size (n = 37, r2= 0.6935, p < 0.001) for
Allometric formula relationships between body size
sockeye salmon are described by; F = 0.2280 L1495L and breeding characters
and E = 0.2289
where L , F, and E are fork
The fecundity and egg size on body size regres~-4-

KAERLYAh4A ET AL. - REPRODUCTIVECHARACTERS OF SOCKEYE AND KOKANEE SALMON

Table 2. Fork length, gonad somatic index (GSI), fecundity, and egg size of Lake Shikotsu kokanee and Bibi River sockeye salmon. Average,
standard errors, and number of individuals are presented as Ave, Se, and n, respectively. Probability level of significance between
populations is represented by P .
Kokanee

Sockeve salmon

salmonage-1.1

age-1.2

249

410

498

Fork length

Ave

(m)

Se

2.7

2.8

5

n

70

16

21

Ave

16.8

17.4

Se
n

0.32
70

0.93
9

17.6
0.68
21

Fecundity

Ave

430

1875

2477

(eggs female^')

Se
n

15.2
70

57.8
16

103
21

Egg size

Ave

5.0

5.0

(m)

Se
n

0.02

0.05
16

5.6
0.04
21

GSI (R)70

sions are frequently calculated as allometric formula(Holtby and Healey, 1986; Beacham et al., 1988).
The allometric regressions by each population (kokanee salmon, age- 1.1, and age-1.2 sockeye salmon)
were observed in relationship between fork length andfecundity (r2> 0.18, P < 0.05j, but were not observedin relationships between fork length and egg size (r2<0.06, P > 0.05),and between fecundity and egg size (r2<0.003, P > 0.05; Table 3). Allometric regressions for
all populations combined were observed in relationships between fork length and fecundity (r? = 0.947, P< 0.001), between fork length and egg size (r2= 0.315,
P < 0.001), and between fecundity and egg size (? =0.508, P < 0.001; Table 3).

Discussion
There appears to be a trade-off between fecundityand egg size in fish (Fleming and Gross, 1990;
Beacham and Murray, 1993). Since Pacific salmon
are semelparous, gametic effort should only reflect
trade-offs with present reproductive demands. These
include competition for breeding sites, parental care,and migration costs (Fleming and Gross, 1990). Vanden Berghe and Gross ( 1 989) found that body size
contributed to adult female fitness in three ways; 1 ) an
increased initial biomass of egg production, 2) theability to acquire a hgh-quality territory for egg devel--5-P< 0.001> 0.05< 0.001< 0.001

opment, and 3) success in nest defense. Fleming and
Gross (1990) reported that egg size of coho salmon
was larger in hatchery populations than in wild ones
because hatchery-released salmon has no breeding
competition in the spawning site. Hatchery-released
Pacific salmon would be useful to evaluate their
gametic effort affected by migration costs because of
exclusion from the breeding competition and parentalcare.
In our analysis for hatchery-released salmon in
Lake Shikotsu and the Bibi River, larger adult female
had higher fecundity, although the GSI showed a constant value (about 17%) in all females from both populations. The fecundity of Lake Shikotsu kokanee
salmon annually fluctuated with change in body size,
although their egg size did not show annual changes.
Fecundity-length relationships for these salmon wereobserved to fit allometry formula. Salmon populations
that allocate a greater proportion of their energy
reserves to migration accomplish their life history by
reducing the energy allocated to gonadal development(Beacham and Murray, 1993). A relative growth coefficient (2.4399) of allometry between fork length andfecundity in Lake Shikotsu kokanee salmon was distinctly higher than that (1.4951) in Bibi River sockeye
salmon. In sockeye salmon, generally, age-.2 femalesconsistently had a higher fecundity per a given bodyS b f . St&%%

SC1. REP. HOKKAIDO SALMONHATCHERY. NO. 49,1995

A

4,000

I
I

0

3,500

*

3,000
v

>

CI

*

0

2,500
3

0

9 2,000
1,500

1

1,000

350

d*

0

n = 37,P = 0.5520, P c 0.001

0

I

I

I

I

400

450

500

550

600

Fork length (L, mm)
6.0 I

5.8
h

5.6

B

I

1

5.0
W

4.8

1

4.6
350

n = 37,P = 0.6935, P c 0.001
I

400

I

500
Fork length (L, mm)
450

I

550

600

Fig. 3. Allometnc relationships between fork length and fecundity (A), and between fork length and egg diameter(B) of age-1.1 and age-1.2 sockeye salmon in the Bibi River dunng the 1990-1993 spawning seasons.

effort (GSI) without breeding competition and parental
care, therefore, fecundity differences within a population may be largely attributable to variation in female
body size caused by environmental variations and perhaps by intraspecific competition such as density

weight than age-.3 fish (Burgner, 1991). Burgner(1991) reported the primary criteria determining
fecundity within populations are size of females andthe number of years spent in the ocean. For hatcheryreleased sockeye salmon having a constant gametic-6-

KAERIYAMA ET AL. - REPRODUCTIVE CHARACTERS OF SOCKEYE AND KOKANEE SALMON

Table 3. Allometric formulas (Y= a Xb)concerning breeding characters of Lake Shikotsu kokanee and Bibi River sockeye salmon. Initial
growth constant and relative growth coefficient are presented by a and b in allometry, respectively. Coefficient of determination isshowed by rz. Probability that b=O is represented by P (NS: P > 0.05, *: P < 0.05. **: P < 0.01, ***: P < 0.001).
Population
Age
Number of individuals

Kokanee3.0
70

All populatlons
combined
107

Sockeye
1.1
16

1.2
21
~

Fork length - Fecundityab

0.00059

2 x 10-1

0.044160.000173.78223

1.75804

2.664 33

r2

2.43995
0.64627

0.632160.18825**

0.94719

P

***

*

***

1.02468
0.262422.319192.756170.140390.10712

Fork length - Egg sizea4.93803b0.00188120.00002NS

0.030960.054080.31537

NS

NS

***

5.01134

4.35479

4.70905

4.026 I6

-0.00073
0.00002NS

0.01752
0.00312NS

0.02098

0.035650.01983NS

0.50849

P
Fecundity - Egg size
a
b
r2

P

***

variable environments. Thus, we think that a range of
optimal egg sizes varies with the habitat and life history pattern of sockeye salmon. The egg size may beaffected by drastic changes in habitable environment
and life history pattern such as regime shift of environment or ocean migration patterns in anadromous
form of the species.
In our conclusion, for sockeye salmon, although
fecundity varies with body size according to environmental factor, egg size may be affected by both environmental and genetic components such as polygene
model within a cohort or population. For hatcheryreleased sockeye salmon, especially, egg size maystrongly relate to genetic component.

effects on growth.
Our results show no relationship between fork
length and egg size in Lake Shikotsu kokanee salmon.
Although age-1.1 sockeye salmon had mean fork
lengths approximately 65% larger than did kokanee
salmon, there was no significant difference in egg size
between these two populations. Age-1.1 female sockeye salmon returning after one year in the ocean are
scarce with small body as termed “jacks” (Burgner,
199 1). Thus, the age- I . I females may not abapt toocean life history. In this study, body size influenced
fecundity but not egg size within kokanee salmon andage-1.1 sockeye salmon populations.
Age-1.2 sockeye salmon had eggs about 12% larger than did kokanee salmon and age-1.1 sockeye
salmon. In salmonids, generally, larger females produce more eggs, and also produce larger eggs (Sargent
et al., 1987). Interaction between fecundity and egg
size has been considered as a trade-off for maximizing
a female’s fitness returns per unit of ovarian resource

Acknowledgments
We thank Dr. Terry D. Beacham of the PacificBiological Station, Dr. William R. Heard of Auke
Bay Fisheries Laboratory, NOAA, NMFS for improving an early draft of the manuscript. This work was(Fleming and Gross, 1990: Beacham and Murray,

supported in a part by a Grant-in-Aid (Bio Cosmos

1993). However, McGinley et al. (1987) reported that
this trade-off model (Smith and Fretwell, 1974) was
not always related with the effect of environmental
variation on offspring investment, and that the variability in offspring size was viewed as an adaptation to

Program) from the Ministry of Agriculture, Forestry
and Fisheries.

References
Bams, R. A. (1976): Survival and propensity for hom-1-

3 l f . %T-L@fB
SCT. REP. HOKKAIDO SALMON HATCHERY, NO. 49,1995

ing as affected by the presence or absence of locally
adapted paternal genes in two transplanted populations of pink salmon (Oncorhynchus gorbuscha). J.
Fish. Rex Bd. Can., 33, 2716-2725.
Beacham, T. D., and C. B. Murray (1986):
Comparative developmental biology of chumsalmon (Oncorhynchus keta) from the Fraser River,
British Columbia. Can. J. Fish. Aquat. Sci., 43,
252-262.
Beacham, T. D., and C. B. Murray (1987): Adaptive
variation in body size, age, morphology, egg size,and developmental biology of chum salmon(Oncorhynchus keta) in British Columbia. Can. J.
Fish. Aquat. Sci., 44, 244-261.
Beacham, T. D., and C. B. Murray (1988): V i a t i o n
in body size, morphology, egg size, and biochemical genetics of pink salmon in British Columbia.
Trans. Am. Fish. SOC., 117, 109-126.
Beacham, T. D., and C. B. Murray (1993): Fecundityand egg size variation in North American Pacificsalmon (Oncorhynchus). J. Fish Biol., 42, 485-508.
Beacham, T. D., C. B. Murray, and R. E. Withler(1988): Age, morphology, developmental biology,
and biochemical genetic variation of Yukon River
fall chum salmon, Oncorhynchus keta, and comparisons with British Columbia populations. Fish.
Bull., 86, 663-674.
Burgner, R. L. ( 1 991): Life history of sockeye salmon(Oncorhynchus nerka). In Pacific Salmon Life
Histories (edited by C. Groot and L. Margolis).
UBC Press, Vancouver. pp. 1-118.
Fleming, I. A., and M. R. Gross (1990): Latitudinalclines: a trade-off between egg number and size in
Pacific salmon. Ecology, 71, 1-11.
Foerster, R. E. (1968): The sockeye salmon,
Oncorhynchus nerka. Bull. Fish. Res. Board Can.,
162,422 p.
Holtby, L. B., and M. C. Healey (1986): Selection for
adult size in female coho salmon (Oncorhynchus
kisutch). Can. J. Fish. Aquat. Sci., 43, 1946-1959.
Kaeriyama, M. (1989): Aspects of salmon ranching inJapan. Physiol. Ecol. Japan, Spec. Volume 1, 625638.
Kaeriyama, M. (1991): Dynamics of the lacustrine

sockeye salmon population in Lake Shikotsu,
Hokkaido. Sci. Rep. Hokkaido Salmon Hatchery,(45), 1-24. (In Japanese with English summary.)
Kaeriyama, M. (1992): Efficient techniques for producing sockeye salmon smolt and improving adultreturns from kokanee salmon. Special Publicationof the National Research Institute of Far Seus
Fisheries, (20), 75.
Kaeriyama, M., S. Urawa, and T. Suzuki (1992):
Anadromous sockeye salmon (Oncorhynchus
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Sci. Rep. Hokkaido Fish Hatchery, (lo), 7-20. (InJapanese with English summary.)

8-

KAERIYAMAET AL. ‑ REPRODUCTIVE

CHARACTERS

体AND

KOKANEE

SALMONI l 歳魚は，体サイズの変化に関わらず，―定の卵サイ

ふ化場産べ二ザケの サイズ，学卵数および卵サイズに関する変異性

OF SOCKEYE・方， べ̲̲ ザケI． 2 歳魚 、ズ (卵径約5 ． Onim) を示した．―の卵サイズのみがと
メ マスやべ二ザケ
I． I歳魚に比べて大型 (卵径約 5． 6mnm) であった． ふ化場産べ.̲ ザケについては，産卵場における繁殖競争とparentalcare カ翔 F除されていることから， 半卵数は生育環境の変化に伴う体サイズの関数で表されるが，卵サイズの変異は量的遺伝モデルのように生育環境要因と遺伝的要因の両構成要素
により支配されている可能性の大きいこと
カ 三示唆された．

帰山雅秀 ・浦和茂彦 ・福音雅章
ふ化場から放流された
支筋湖産ヒメマスとそれを起源とする美々川放流べ̲ ザケの親佳の体サイズ，羊卵数および
卵 サイズに関する
変異性が調べられた．各々の個体群において，卒り口数と体サイズは連動して変化し 両者の間にはallometry式が適合した．ヒメマスとべ二ザケ―

9―