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Hybridization effects of three populations of Chlamys farreri from China, Korea and Japan.


ABSTRACT Populations of Chlamys farreri are distributed along the coasts of the northern seacoast of China, the Korean Peninsula and the Pacific seacoast, Japan. These populations represent potentially different genetic resources in their production traits important to aquaculture aquaculture, the raising and harvesting of fresh- and saltwater plants and animals. The most economically important form of aquaculture is fish farming, an industry that accounts for an ever increasing share of world fisheries production.  and for their contribution to hybridization hybridization /hy·brid·iza·tion/ (hi?brid-i-za´shun)
1. crossbreeding; the act or process of producing hybrids.

2. molecular hybridization

3.
 breeding programs A breeding program is the planned breeding of a group of animals or plants, usually involving at least several individuals and extending over several generations. Breeding programs are commonly employed in several fields where humans wish to manage the characteristics of their . To determine their characteristics, crosses between and within three populations of Zhikong scallop scallop or pecten, marine bivalve mollusk. Like its close relative the oyster, the scallop has no siphons, the mantle being completely open, but it differs from other mollusks in that both mantle edges have a row of steely blue "eyes" and . Chlamys farreri, were made (Chinese culture population [CC] female x Japanese wild population [JW] male, Japanese wild population (JW) female x Chinese culture population [CC] male, Chinese culture population [CC] female x Korean wild population [KW] male, Japanese wild population [JW] female x Korean wild population [KW] male) and the offspring reared to 24 months of age. Nine consecutive measurements, approximately 60 days apart, were recorded on wet body weight and shell length, width and height. The phenotypic phe·no·type  
n.
1.
a. The observable physical or biochemical characteristics of an organism, as determined by both genetic makeup and environmental influences.

b.
 and heterotic response of each sample for growth and survival were compared. The results indicated that there existed positive though varying degrees of heterosis heterosis (hĕt'ərō`sĭs): see hybrid.
heterosis
 or hybrid vigor

Increase in such characteristics as size, growth rate, fertility, and yield of a hybrid organism over those of its parents.
 for growth in wet body weight, shell length, shell height, shell width, and survival. Also, there were measurable differences among the reciprocal crosses Noun 1. reciprocal cross - hybridization involving a pair of crosses that reverse the sexes associated with each genotype
reciprocal

hybridisation, hybridization, hybridizing, interbreeding, crossbreeding, crossing, cross - (genetics) the act of mixing
 in their performance traits. The results of this study suggest that hybridization is an important tool to increase the performance and fitness (survival) of cultured scallops.

KEY WORDS: Chlamys farreri, hybrid, geographic population, growth, heterosis, hybrid vigor hybrid vigor
n.
Increased vigor or other superior qualities arising from the crossbreeding of genetically different plants or animals. Also called heterosis.
 

INTRODUCTION

Chlamys farreri (Jones et Preston), Chlamys Roding, Pectinidae, Pterioidae, Bivalvia, Mollusca is distributed along the coast of Northern China, the coast of Southern Korea and Northern Korea, the coast of Japan, and the coast of Eastern Russia Eastern Russia is the region of Russia between the Ural Mountains and the Pacific Ocean.
  • Siberia
  • Russian Far East
, it is also widely distributed Adj. 1. widely distributed - growing or occurring in many parts of the world; "a cosmopolitan herb"; "cosmopolitan in distribution"
cosmopolitan

bionomics, environmental science, ecology - the branch of biology concerned with the relations between organisms
 around the sea area of Shandong province and Liaoning province in China (Pan et al. 2002). The first batch of seedlings were triumphantly cultivated in Dalian, Liaoning province in 1974, and intensively cultured in China where the scallop industry has grown rapidly over the last 20 years. The cultured area has reached 400,000 [mu] and production yield of 919,591 tons; tiptop yield reached more than one million tons, and Chlamys farreri culture is one of the three mainstay mariculture mariculture

marine aquaculture.
 industries in China, which is now one of the top producers of cultured scallop in the world (Xiang 2002). Chlamys farreris grow relatively slow (1.5-2 y to market size of 6-8 cm). The primary problem of Chlamys farreri culture is cosmically cos·mic   also cos·mi·cal
adj.
1. Of or relating to the universe, especially as distinct from Earth.

2. Infinitely or inconceivably extended; vast:
 death because of the decline of the idioplasm and disease resistance (Wang et al. 2002). To overcome the obstacle, the new breeding method, especially crossbreeding crossbreeding /cross·breed·ing/ (-bred-ing) hybridization; the mating of organisms of different strains or species.

crossbreeding

hybridization; the mating of organisms of different strains or species, e.g.
 should be applied.

Technologic developments have made hatchery hatchery

a commercial establishment dedicated to the hatching of bird eggs to provide day old chicks and poults to the poultry industry.


hatchery liquid
the contents of unfertilized eggs. Used in petfood manufacture.
 production of scallop's feasible (Castagna & Duggan 1971, Gruffydd & Beaumont 1972). The scallop species, Chlamys farreri, has become an important aquaculture species along the coast of northern China with the first hatchery production of seedling scallops in the 1970s (Wang 1981). However, seedling quality (idioplasm) degeneration degeneration /de·gen·er·a·tion/ (de-jen?er-a´shun) deterioration; change from a higher to a lower form, especially change of tissue to a lower or less functionally active form.  and fitness depression have become major technical obstacles to intensive hatchery production of scallops.

Inter-species and intra-specific hybridization has received attention front researchers in aquaculture because it earl improve productivity through hybrid vigor and combine desirable characteristics found in one species with those of another (or in one population with those of another). Experimental attempts to produce inter-species hybrids in the oyster oyster, edible bivalve mollusk found in beds in shallow, warm waters of all oceans. The shell is made up of two valves, the upper one flat and the lower convex, with variable outlines and a rough outer surface. , the scallop, and the hard clam have reported few successes. It has been reported that spawn To launch another program from the current program. The child program is spawned from the parent program.

(operating system) spawn - To create a child process in a multitasking operating system. E.g.
 rate, hatch rate, and the survival of hybrid progeny PROGENY - 1961. Report generator for UNIVAX SS90.  to post-larval stages are low compared with the intra-specific matings and then some inter-species hybrids are sterile sterile /ster·ile/ (ster´il)
1. unable to produce offspring.

2. aseptic.


ster·ile
adj.
1. Not producing or incapable of producing offspring.

2.
 (Hedgecock 1987, Longwell 1987, Menzel 1987, Xiang & Chen 1991, Ouang & Liu 1959, Clark & Talbot 1973, Ke & Tian Tian
 or T'ien
(Chinese; “Heaven”)

In indigenous Chinese religion, the supreme power reigning over humans and lesser gods. The term refers to a deity, to impersonal nature, or to both.
 2000, Leighton & Lewis 1982, Zhou & Gao 1982).

Intra specific hybrids can produce heterosis and combine desirable characteristics to improve productivity because different populations represent potentially different genetic resources (Manzi & Hadley 1991, Newkirk & Waugh 1977, Newkirk 1978, Mallet mallet,
n a hammering instrument.

mallet, hard,
n a small hammer with a leather-, rubber-, fiber-, or metal-faced head; used to supply force or to supplement hand force for the compaction of foil or amalgam and to seat cast
 & Haley 1983, Stiles Stiles can refer to: People
  • Bert Stiles, short story writer
  • Charles Wardell Stiles, American zoologist
  • Edgar Stiles, character on the popular drama 24
  • Ezra Stiles, president of Yale College
  • Innis Stiles, singer, musician
 1978, Bower & Blackbourn 1997, Cruz & Ibarra 1997, Chevassus 1983).

The purpose of this study is to investigate population differences and heterosis in growth traits and survival among three samples taken from Chinese, Japanese and Korean populations of the Zhikong scallop, Chlamys farreri. The population samples were mated to produce pure and reciprocal cross progeny.

MATERIALS AND METHODS

Collection and Maintenance

The sample of the Chinese scallop (CC) was collected from the cultured population in Dalian Bay Dalian Bay (also spelled Talien Bay) is a roughly rectangular arm of the Korea Bay, oriented long side parallel to the land due south-east of the modern port city of Dalian, and serving as the roadstead for the harbor. , Dalian City, Liaoning province (north latitude latitude, angular distance of any point on the surface of the earth north or south of the equator. The equator is latitude 0°, and the North Pole and South Pole are latitudes 90°N and 90°S, respectively.  38[degrees]42'N, east longitude longitude (lŏn`jĭtd'), angular distance on the earth's surface measured along any latitude line such as the equator east or west of the prime meridian.  121[degrees]31'E, and annual temperature range 1.0-22.8[degrees]C). The sample of the Korean scallop (KW) was collected from a wild population on the sea coast of Korea (north latitude 35[degrees]28'N, east longitude 130[degrees]28'E, and annual temperature range 4.5-23.7[degrees]C). The sample of the Japanese scallop Japanese scallop

see pecten yessoensis.
 (JW), also from a wild population, was collected from the Pacific seacoast of Japan (north latitude 40[degrees]52'N, east longitude 142[degrees]22'E, annual temperature range 2.5-21.6[degrees]C. Each sample was held for half year in a maturation maturation /mat·u·ra·tion/ (mach-u-ra´shun)
1. the process of becoming mature.

2. attainment of emotional and intellectual maturity.

3.
 unit prior to mating. The diet consisted of fresh Isochrysis zhanjiangensis and Platymonas sp.

Experimental Design and Cross-fertilization

Fertilization fertilization, in biology, process in the reproduction of both plants and animals, involving the union of two unlike sex cells (gametes), the sperm and the ovum, followed by the joining of their nuclei.  between and within the three population samples were conducted at room temperature (19-21 [degrees]C): CC male x CC female, JW male x CC female, KW male x CC female, CC male x JW female, JW male x JW female, KW male x JW female. Only male scallops were included in the sample taken from the KW population. Thus, the within and cross genotype genotype (jēn`ətīp'): see genetics.
genotype

Genetic makeup of an organism. The genotype determines the hereditary potentials and limitations of an individual.
 fertilizations involving KW females are not included in this study because of no female spawning eggs.

A random sample of 80-100 adult Chlamys faarreri individuals (7-10 cm length) from each of the three populations were taken for spawning. Each sample was dried in the shade for 2 hours and placed in a separate container filled with UV-sterilized filtered seawater seawater

Water that makes up the oceans and seas. Seawater is a complex mixture of 96.5% water, 2.5% salts, and small amounts of other substances. Much of the world's magnesium is recovered from seawater, as are large quantities of bromine.
 to collect sperm sperm or spermatozoon (spûr'mətəzō`ən, –zō`ŏn), in biology, the male gamete (sex cell), corresponding to the female ovum in organisms that reproduce sexually.  and ovum. Fertilization was accomplished through mixing of the diluted di·lute  
tr.v. di·lut·ed, di·lut·ing, di·lutes
1. To make thinner or less concentrated by adding a liquid such as water.

2. To lessen the force, strength, purity, or brilliance of, especially by admixture.
 sperm and diluted ovum containers. After 2-4 hours the sperm-ovum mixture was rinsed in seawater two to three times and incubated in seawater in 100 L plastic tanks at room temperature.

Larval larval

1. pertaining to larvae.

2. larvate.


larval migrans
see cutaneous and visceral larva migrans.
 and Juvenile Rearing

Larvae Larvae, in Roman religion
Larvae: see lemures.
 were reared on a diet of mixed algae algae (ăl`jē) [plural of Lat. alga=seaweed], a large and diverse group of primarily aquatic plantlike organisms. These organisms were previously classified as a primitive subkingdom of the plant kingdom, the thallophytes (plants that , Isochrysis zhanjiangensis and Platymonas sp. All cultures were carried out in filtered seawater (FSW FSW Friction Stir Welding
FSW Flight Software
FSW Full Spectrum Warrior (video game)
FSW Family Support Worker
FSW Female Sex Worker
FSW Fox Sports World (cable TV channel) 
) at 19-23[degrees]C, the approximate ambient Surrounding. For example, ambient temperature and humidity are atmospheric conditions that exist at the moment. See ambient lighting.  sea water temperature. Each day 50% to 75% of the culture water was removed by reverse filtration/siphoning and replaced with fresh FSW. Fresh air was continuously pumped into each culture.

Culture of Adults

After 2 months of age the cultured juveniles from each group were placed separately into plastic cages and suspended in the sea. Body weight and shell height and width and length measurements were recorded every 2-3 months, when the cages were changed, until 24 months of age.

Measurement Date

Although the experimental design called for weight and linear measurements to be recorded at 60-day intervals, weather conditions and workload sometimes precluded doing this.

Measurement and Statistical Analyses

From each progeny subclass In programming, to add custom processing to an existing function or subroutine by hooking into the routine at a predefined point and adding additional lines of code.

subclass - derived class
 100-150 offspring were sampled for measurements of body size (shell length, S-L S-L Personnel/Administrative Staff Section ; shell height, S-H; shell width, S W) and body weight (B-W) at approximately 2-month intervals. Within each test period the percent heterosis (H) of each cross was computed as:

(1) H(%) = [F.sub.1] - 1/2 [([P.sub.1] + [P.sub.2])] / 1/2 [([P.sub.1] + [P.sub.2])] x 100

where, [F.sub.1], [P.sub.1], [P.sub.2] represent trait trait (trat)
1. any genetically determined characteristic; also, the condition prevailing in the heterozygous state of a recessive disorder, as the sickle cell trait.

2. a distinctive behavior pattern.
 average of the first generation cross, parent l and parent 2, respectively. Incremental Additional or increased growth, bulk, quantity, number, or value; enlarged.

Incremental cost is additional or increased cost of an item or service apart from its actual cost.
 change (G) in body weight and in each linear measurement for each period was computed as:

(2) G = ([W.sub.2] - [W.sub.1]) / ([t.sub.2] - [t.sub.1]),

where [t.sub.1] and [t.sub.2] represent time at the beginning and end of each period and [W.sub.1] and [W.sub.1] represent the menstruation menstruation, periodic flow of blood and cells from the lining of the uterus in humans and most other primates, occurring about every 28 days in women. Menstruation commences at puberty (usually between age 10 and 17).  value at the beginning and end of each period.

RESULTS

Comparison of Heterosis of Different Crossbred crossbred

progeny of a mating between two animals which are purebreds of different breeds, e.g. crossbred sheep are usually offspring of matings between merinos and British breeds.
 for Different Characters in Each Period

The average and standard deviation In statistics, the average amount a number varies from the average number in a series of numbers.

(statistics) standard deviation - (SD) A measure of the range of values in a set of numbers.
 of each progeny group for body weight and linear measurement, including calculated heterosis for the reciprocal crossbred samples CC x JW and JW x CC, for each test period are included in Table 1. The Chinese cultured population has similar body weight growth, as does the Japanese wild population. Across the nine measurement intervals the [F.sub.1] hybrid progeny of three Chlamys faarreri samples exhibited a wide range of heterosis, from 4% to 50% (see Table 1). Among the [F.sub.1] progeny the JW x KW cross exhibited the most heterosis followed by the CC x KW, JW x CC, and CC x JW crosses. Among the growth traits there were distinct differences in the magnitude of heterosis expression. Shell height tended to be more heterotic than shell length followed by shell width and wet weight. The expression of heterosis in each trait tended to decrease as age of the scallop increased.

In this study the [F.sub.1] hybrid combinations exhibited satisfactory levels of heterosis over the nine evaluation periods Evaluation period

The time interval over which funds assess a money manager's performance.
. The JW female x KW male and CC female x KW male F1 were the optimal crosses, than can be applied in aquaculture production.

Comparison of the Growth Rate for Four Hybrids and the Purebred purebred

progeny derived from at least several generations of animals of the same breed.


purebred herds
herds (or flocks) composed of purebred animals. Not necessarily registered animals. Distinct from crossbred herds.
 

The means ([delta], ccm/d, and mg/d) for growth rate of shell length, width, and height and for body weight by age in months for the CC and JW parental samples and four F1 progeny samples are included in Figure 1.

[FIGURE 1 OMITTED]

Growth rate for each trait in each sample reached its pinnacle pinnacle (pĭn`ĭkəl), minor architectural motif of vertical tapering shape, usually crowning a pier, buttress, or gable. Although sometimes it appears in Renaissance design, as in the Certosa di Pavia, it is almost exclusively a medieval  at 13 months of age. The incremental changes in growth were greater for each trait in each hybrid sample than in the pure parental samples.

The Growth Trend of Chlamys Farreri for Different Characters for Four Hybrids and the Offspring of Pure Parents by Age in Months

Based on the serial measurements Serial measurements
A series of measurements looking for an increase or decrease over time.

Mentioned in: Tumor Markers
 of wet weight and shell length increase in parental and [F.sub.1] sample of Chlamys farreri, growth differences among the parental and [F.sub.1] progeny samples were evaluated (Fig. 2).

[FIGURE 2 OMITTED]

The growth in shell length tends to plateau plateau, elevated, level or nearly level portion of the earth's surface, larger in summit area than a mountain and bounded on at least one side by steep slopes, occurring on land or in oceans.  at approximately 16 months of age. The growth curve for both wet weight and shell length was greater in the [F.sub.1] progeny samples than in the pure parental samples. Among the [F.sub.1] progeny samples, the growth curve for wet weight and shell length tended to be different.

Comparison of Growth at 24 Months of Age for Three Hybrids and the Offspring of Their Parental Populations

At 24 months of age the wet weight of the [F.sub.1] progeny samples was heavier than their pure parental samples (see Fig. 2 A). Much of the difference between the [F.sub.1] progeny and their parental pure counterparts can be attributed to heterosis. Wet weight is the most important trait in scallop production and hybridization can be a useful tool in improving wet weight growth. However, not all [F.sub.1] combinations result in equal growth in wet weight. Therefore, experimental evaluation of crosses between sampled populations becomes a necessary precondition pre·con·di·tion  
n.
A condition that must exist or be established before something can occur or be considered; a prerequisite.

tr.v.
 to include hybridization of populations in commercial scallop culturing. Figure 2 B includes the differences at 24 months of age among the [F.sub.1] and parental pure progeny samples for shell length. The shell length of the F1 samples was much greater than those of the parental populations. As was the case for wet weight, heterosis was an important part of the increase in shell length in the [F.sub.1] progeny samples.

DISCUSSION

Comparison of the Survival for Four Hybrids and the Offspring of Two Parent Populations

The percentage of survival from the zygotes to juveniles of 2 months of age for four hybrids and the offspring of two parent populations were calculated in Table 1. The percentage of survival for four hybrids were significantly greater than the purebred (P < 0.05), and the percentage of survival for Chinese purebred was significantly greater than that of Japanese purebred, there were no significant differences of survival percentage among the four (P > 0.05).

Comparison of the Growth for Three Hybrids and the Offspring of Three Parent Populations

Our evaluation of tour phenotypic traits, shell length, shell width, shell height and body weight, suggest genetic differences among two scallop populations and their reciprocal [F.sub.1] crosses evaluated simultaneously under the same environmental conditions. From age 5 months to 24 months, there were differences in body weight and shell linear sizes over the sampling period. The samples from the Chinese cultured population and the Japanese wild population were similar, but crossbred offspring were consistently faster (P < 0.05) growing than purebred offspring (see Table 1).

Comparison of the Growth Among the Different Periods

The growth rates Growth Rates

The compounded annualized rate of growth of a company's revenues, earnings, dividends, or other figures.

Notes:
Remember, historically high growth rates don't always mean a high rate of growth looking into the future.
 for each of the four phenotypic traits included in this study are influenced by seasonal environmental conditions. During winter months the cold water causes the growth rate of scallops to approach zero or even lose body weight and size. However, during the warmer seasons of summer and tall growth rate rapidly increases.

Comparison Among the Growth Rate of Four Phenotypic Traits Over the Evaluation Period

The growth rates of each of the four phenotypic traits, shell length, shell width, shell height, and body weight tend to be different. Among these traits body weight growth is the fastest followed by shell length, shell width, and shell height. Therefore, among the linear shell measurements the shell length should he the most important selection trait in a breeding program.

Breeding and Heterosis Utilization of Chlamys Farreri

Scallop breeding has received considerable research attention because the development of aquaculture has increased. Developments in artificial insemination artificial insemination, technique involving the artificial injection of sperm-containing semen from a male into a female to cause pregnancy. Artificial insemination is often used in animals to multiply the possible offspring of a prized animal and for the breeding  and juvenile rearing techniques have succeeded in improving the efficiency of scallop culturing. However, performance of the economic traits and purity of the cultural stocks have been reduced because of the idioplasm retrogression retrogression /ret·ro·gres·sion/ (ret?ro-gresh´un) degeneration; deterioration; regression; return to an earlier, less complex condition.

ret·ro·gres·sion
n.
1.
 and the parental scallops are wild populations. Thus, it is difficult to expect stable levels of heterosis. Selective breeding
This article focuses on selective breeding in domesticated animals. For alternate uses, see artificial selection.


Selective breeding in domesticated animals is the process of developing a cultivated breed over time.
 and evaluation of new breeds and strains are the bases of the hybridization breeding programs. Such breeding programs can provide pure hybridization parents and overcome the idioplasm retrogression. The research reported here shows that for wet shell weight and linear shell measurements in crosses among cultured and wild populations of the Chlamys farreri scallop, positive heterosis are expressed at levels sufficient to warrant systematic hybridization in the culture of scallops.
TABLE 1.
Comparison of survival and heterosis of growth of hybrids with their
parental populations. Values marked with different superscripts are
significantly different from each other at 0.05.

               Combination
               Character/              CC
    Age           Index           X [+ or -] S

Survival (%)                  82.9 (b) [+ or -] 8.9
               S-L/cm         1.35 (b) [+ or -] 0.20
               S-H/cm         1.64 (b) [+ or -] 0.24

5 months       S-W/cm         0.44 (c) [+ or -] 0.05
               B-W/g          1.23 (c) [+ or -] 0.26
               S-L/cm         1.63 (b) [+ or -] 0.18
               S-H/cm         1.99 (c) [+ or -] 0.23

9 months       S-W/cm         0.53 (b) [+ or -] 0.07
               B-W/g          1.49 (c) [+ or -] 0.35
               S-L/cm         1.98 (b) [+ or -] 0.38
               S-H/cm         2.39 (b) [+ or -] 0.44

11 months      S-W/cm         0.68 (b) [+ or -] 0.11
               B-W/g          1.93 (b) [+ or -] 0.52
               S-L/cm             3.80 [+ or -] 0.39
               S-H/cm             4.33 [+ or -] 0.36

13 months      S-W/cm         1.34 (b) [+ or -] 0.15
               B-W/g         12.55 (c) [+ or -] 2.08
               S-L/cm         5.35 (c) [+ or -] 0.38
               S-H/cm         5.60 (b) [+ or -] 0.42

16 months      S-W/cm         2.00 (b) [+ or -] 0.20
               B-W/g         30.63 (b) [+ or -] 4.81
               S-L/cm         5.39 (b) [+ or -] 0.40
               S-H/cm         5.65 (b) [+ or -] 0.38

18 months      S-W/cm             2.01 [+ or -] 0.20
               B-W/g         27.70 (c) [+ or -] 6.96
               S-L/cm         5.47 (b) [+ or -] 0.45
               S-H/cm         5.79 (b) [+ or -] 0.43

20 months      S-W/cm             2.05 [+ or -] 0.23
               B-W/g         30.98 (b) [+ or -] 9.50
               S-L/cm         5.54 (b) [+ or -] 0.49
               S-H/cm         6.10 (b) [+ or -] 0.48

22 months      S-W/cm             2.08 [+ or -] 0.21
               B-W/g         36.15 (c) [+ or -] 8.03
               S-L/cm         5.81 (b) [+ or -] 0.53

24 months      S-H/cm         6.42 (b) [+ or -] 0.54
               S-W/cm         2.24 (b) [+ or -] 0.20
               B-W/g         39.32 (c) [+ or -] 8.32

                                       [F.sub.1] (CC x JW)

               Combination
               Character/                                 Heterosis
    Age           Index             X [+ or -] S             (%)

Survival (%)                    90.0 (a) [+ or -] 10.2      13.31
               S-L/cm           1.75 (a) [+ or -] 0.22      30.32
               S-H/cm           2.16 (a) [+ or -] 0.290     27.86

5 months       S-W/cm           0.53 (b) [+ or -] 0.09      18.73
               B-W/g            1.54 (b) [+ or -] 0.43      12.06
               S-L/cm           2.12 (a) [+ or -] 0.42      30.36
               S-H/cm           2.42 (b) [+ or -] 0.37      23.05

9 months       S-W/cm           0.67 (a) [+ or -] 0.11      23.53
               B-W/g            1.91 (b) [+ or -] 0.79      28.37
               S-L/cm           2.39 (a) [+ or -] 0.45      18.43
               S-H/cm           2.85 (a) [+ or -] 0.60      18.77

11 months      S-W/cm           0.79 (a) [+ or -] 0.13      14.31
               B-W/g            2.85 (a) [+ or -] 0.53      44.36
               S-L/cm               4.29 [+ or -] 0.52      10.51
               S-H/cm               4.66 [+ or -] 0.57       7.12

13 months      S-W/cm           1.56 (a) [+ or -] 0.16      14.01
               B-W/g           16.15 (b) [+ or -] 4.03      25.87
               S-L/cm           5.76 (b) [+ or -] 0.57       8.10
               S-H/cm           6.28 (a) [+ or -] 0.59      11.56

16 months      S-W/cm        2.15 (a)(b) [+ or -] 0.20       6.90
               B-W/g           38.34 (a) [+ or -] 9.26      22.49
               S-L/cm           5.72 (b) [+ or -] 0.48       5.52
               S-H/cm           6.23 (a) [+ or -] 0.51       8.73

18 months      S-W/cm               2.23 [+ or -] 024        8.54
               B-W/g           34.83 (b) [+ or -] 8.21      20.08
               S-L/cm           5.92 (a) [+ or -] 0.66       7.37
               S-H/cm           6.40 (a) [+ or -] 0.70      10.08

20 months      S-W/cm               2.23 [+ or -] 0.56       7.14
               B-W/g           38.40 (a) [+ or -] 19.50     23.43
               S-L/cm           6.00 (a) [+ or -] 0.52       6.40
               S-H/cm           6.51 (a) [+ or -] 0.52       6.67

22 months      S-W/cm               2.22 [+ or -] 0.19       4.59
               B-W/g           40.40 (b) [+ or -] 9.50      10.52
               S-L/cm           6.14 (a) [+ or -] 0.47       4.93

24 months      S-H/cm           6.69 (a) [+ or -] 0.46       4.84
               S-W/cm           2.41 (a) [+ or -] 0.31       7.32
               B-W/g            44.8 (b) [+ or -] 9.98      12.15

                                        [F.sub.1] (JW x CC)

               Combination
               Character/                                  Heterosis
    Age           Index             X [+ or -] S              (%)

Survival (%)                     87.6 (a) [+ or -] 9.6       10.28
               S-L/cm            1.81 (a) [+ or -] 0.27      34.55
               S-H/cm            2.22 (a) [+ or -] 0.27      31.95

5 months       S-W/cm            0.57 (a) [+ or -] 0.07      27.71
               B-W/g             1.68 (a) [+ or -] 0.59      21.86
               S-L/cm            2.18 (a) [+ or -] 0.23      34.09
               S-H/cm         2.56 (a)(b) [+ or -] 0.24      29.95

9 months       S-W/cm            0.71 (a) [+ or -] 0.07      29.60
               B-W/g             2.15 (a) [+ or -] 0.61      44.73
               S-L/cm            2.42 (a) [+ or -] 0.40      19.46
               S-H/cm            2.88 (a) [+ or -] 0.47      19.90

11 months      S-W/cm            0.82 (a) [+ or -] 0.12      18.28
               B-W/g             2.97 (a) [+ or -] 0.53      50.80
               S-L/cm                4.53 [+ or -] 0.22      16.78
               S-H/cm                5.00 [+ or -] 0.352     15.03

13 months      S-W/cm            1.62 (a) [+ or -] 0.11      18.25
               B-W/g         17.45 (a)(b) [+ or -] 1.68      36.00
               S-L/cm            5.94 (a) [+ or -] 0.40      11.48
               S-H/cm            6.44 (a) [+ or -] 0.38      14.40

16 months      S-W/cm            2.21 (a) [+ or -] 0.15       9.60
               B-W/g            39.94 (a) [+ or -] 6.60      27.57
               S-L/cm            6.01 (a) [+ or -] 0.37      10.93
               S-H/cm            6.48 (a) [+ or -] 0.38      13.17

18 months      S-W/cm                2.25 [+ or -] 0.16       9.78
               B-W/g            38.05 (a) [+ or -] 5.85      31.19
               S-L/cm            6.11 (a) [+ or -] 0.51      10.83
               S-H/cm            6.50 (a) [+ or -] 0.55      11.82

20 months      S-W/cm                2.26 [+ or -] 0.19       8.48
               B-W/g            39.59 (a) [+ or -] 8.99      27.17
               S-L/cm            6.15 (a) [+ or -] 0.55       9.17
               S-H/cm            6.53 (a) [+ or -] 0.54       7.06

22 months      S-W/cm                2.30 [+ or -] 0.45       8.40
               B-W/g         41.89 (a)(b) [+ or -] 10.1      14.59
               S-L/cm            6.18 (a) [+ or -] 0.44       5.71

24 months      S-H/cm            6.67 (a) [+ or -] 0.45       4.42
               S-W/cm            2.45 (a) [+ or -] 0.21       9.42
               B-W/g            45.26 (b) [+ or -] 9.25      13.19

               Combination
               Character/               JW
    Age           Index            X [+ or -] S

Survival (%)                  75.9 (c) [+ or -] 23.2
               S-L/cm         1.34 (b) [+ or -] 0.26
               S-H/cm         1.73 (b) [+ or -] 0.31

5 months       S-W/cm         0.45 (c) [+ or -] 0.10
               B-W/g          1.53 (b) [+ or -] 0.40
               S-L/cm         1.62 (b) [+ or -] 0.24
               S-H/cm         1.95 (b) [+ or -] 0.29

9 months       S-W/cm         0.56 (b) [+ or -] 0.17
               B-W/g          1.48 (c) [+ or -] 0.43
               S-L/cm         2.06 (b) [+ or -] 0.39
               S-H/cm         2.41 (b) [+ or -] 0.41

11 months      S-W/cm         0.71 (b) [+ or -] 0.12
               B-W/g          2.01 (b) [+ or -] 0.52
               S-L/cm         3.95 (b) [+ or -] 0.47
               S-H/cm         4.37 (b) [+ or -] 0.42

13 months      S-W/cm         1.40 (b) [+ or -] 0.14
               B-W/g         13.11 (c) [+ or -] 3.02
               S-L/cm         5.32 (b) [+ or -] 0.47
               S-H/cm         5.65 (b) [+ or -] 0.48

16 months      S-W/cm         2.03 (b) [+ or -] 0.20
               B-W/g         31.98 (b) [+ or -] 7.42
               S-L/cm         5.44 (c) [+ or -] 0.48
               S-H/cm         5.81 (b) [+ or -] 0.57

18 months      S-W/cm             2.09 [+ or -] 0.21
               B-W/g         30.31 (c) [+ or -] 7.83
               S-L/cm         5.56 (b) [+ or -] 0.51
               S-H/cm         5.84 (b) [+ or -] 0.76

20 months      S-W/cm             2.11 [+ or -] 0.25
               B-W/g         31.28 (b) [+ or -] 7.59
               S-L/cm         5.73 (b) [+ or -] 0.51
               S-H/cm         6.09 (b) [+ or -] 0.63

22 months      S-W/cm             2.17 [+ or -] 0.22
               B-W/g         36.97 (c) [+ or -] 8.94
               S-L/cm         5.90 (c) [+ or -] 0.52

24 months      S-H/cm         6.34 (b) [+ or -] 0.49
               S-W/cm         2.24 (b) [+ or -] 0.20
               B-W/g         40.65 (c) [+ or -] 10.29

               Combination          [F.sub.1]
               Character/           (JW x KW)
    Age           Index           X [+ or -] S

Survival (%)                  90.8 (a) [+ or -] 11.4
               S-L/cm         1.85 (a) [+ or -] 0.31
               S-H/cm         2.20 (a) [+ or -] 0.28

5 months       S-W/cm         0.59 (a) [+ or -] 0.08
               B-W/g          1.73 (a) [+ or -] 0.47
               S-L/cm         2.30 (a) [+ or -] 0.22
               S-H/cm         2.72 (a) [+ or -] 0.22

9 months       S-W/cm         0.72 (a) [+ or -] 0.08
               B-W/g          2.24 (a) [+ or -] 0.64
               S-L/cm         2.55 (a) [+ or -] 0.40
               S-H/cm         2.96 (a) [+ or -] 0.48

11 months      S-W/cm         0.83 (a) [+ or -] 0.16
               B-W/g          3.05 (a) [+ or -] 0.52
               S-L/cm             4.59 [+ or -] 0.56
               S-H/cm             5.23 [+ or -] 0.54

13 months      S-W/cm         1.66 (a) [+ or -] 0.22
               B-W/g         18.48 (a) [+ or -] 4.83
               S-L/cm         5.92 (a) [+ or -] 0.53
               S-H/cm         6.49 (a) [+ or -] 0.47

16 months      S-W/cm         2.23 (a) [+ or -] 0.19
               B-W/g         40.25 (a) [+ or -] 9.45
               S-L/cm         6.02 (a) [+ or -] 0.43
               S-H/cm         6.51 (a) [+ or -] 0.54

18 months      S-W/cm         2.26 (a) [+ or -] 0.22
               B-W/g         39.67 (a) [+ or -] 0.52
               S-L/cm         6.07 (a) [+ or -] 0.44
               S-H/cm         6.52 (a) [+ or -] 0.56

20 months      S-W/cm             2.26 [+ or -] 0.79
               B-W/g         41.34 (a) [+ or -] 8.72
               S-L/cm         6.13 (a) [+ or -] 0.49
               S-H/cm         6.62 (a) [+ or -] 0.48

22 months      S-W/cm         2.35 (a) [+ or -] 0.65
               B-W/g         43.34 (a) [+ or -] 5.84
               S-L/cm         6.21 (a) [+ or -] 0.51

24 months      S-H/cm         6.71 (a) [+ or -] 0.51
               S-W/cm         2.43 (a) [+ or -] 0.19
               B-W/g         48.48 (a) [+ or -] 9.58

               Combination           [F.sub.1]
               Character/            (CC x KW)
    Age           Index             X [+ or -] S

Survival (%)                     92.1 (a) [+ or -] 13.1
               S-L/cm            1.83 (a) [+ or -] 0.29
               S-H/cm            2.10 (a) [+ or -] 0.24

5 months       S-W/cm            0.57 (a) [+ or -] 0.07
               B-W/g             1.70 (a) [+ or -] 0.42
               S-L/cm            2.23 (a) [+ or -] 0.34
               S-H/cm            2.63 (b) [+ or -] 0.30

9 months       S-W/cm            0.71 (a) [+ or -] 0.08
               B-W/g             2.18 (a) [+ or -] 0.77
               S-L/cm            2.47 (a) [+ or -] 0.39
               S-H/cm            2.86 (a) [+ or -] 0.42

11 months      S-W/cm            0.83 (a) [+ or -] 0.13
               B-W/g             2.97 (a) [+ or -] 0.51
               S-L/cm                4.59 [+ or -] 0.61
               S-H/cm                5.16 [+ or -] 0.54

13 months      S-W/cm            1.65 (a) [+ or -] 0.17
               B-W/g            18.29 (a) [+ or -] 4.38
               S-L/cm            5.89 (a) [+ or -] 0.59
               S-H/cm            6.30 (a) [+ or -] 0.56

16 months      S-W/cm         2.12 (a)(b) [+ or -] 0.17
               B-W/g            37.62 (a) [+ or -] 8.40
               S-L/cm            5.94 (a) [+ or -] 0.56
               S-H/cm            6.38 (a) [+ or -] 0.52

18 months      S-W/cm                2.14 [+ or -] 0.20
               B-W/g          36.8 (a)(b) [+ or -] 0.48
               S-L/cm            5.97 (a) [+ or -] 0.53
               S-H/cm            6.43 (a) [+ or -] 0.46

20 months      S-W/cm                2.15 [+ or -] 0.19
               B-W/g            40.35 (a) [+ or -] 7.06
               S-L/cm            6.10 (a) [+ or -] 0.54
               S-H/cm            6.52 (a) [+ or -] 0.39

22 months      S-W/cm            2.26 (a) [+ or -] 0.17
               B-W/g            42.54 (a) [+ or -] 6.12
               S-L/cm            6.16 (a) [+ or -] 0.48

24 months      S-H/cm            6.65 (a) [+ or -] 0.42
               S-W/cm            2.42 (a) [+ or -] 0.19
               B-W/g         46.49 (a)(b) [+ or -] 7.7


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LIU XIAOLIN, (1,3) CHANG YAQING, (2) XIANG JIANHAI (1), *

(1) Academy of Sciences, Qingdao 266071, China; (2) Key Lab of Marine Ecology Marine ecology

An integrative science that studies the basic structural and functional relationships within and among living populations and their physical-chemical environments in marine ecosystems.
 in Aquaculture of Ministry of Agriculture, Dalian Fisheries University Dalian Fisheries University (大连水产学院) is a university located in Dalian, China. Founded in 1952, it is the sole university featuring fisheries science courses in northern China. Over 7,000 students are enrolled there. , Dalian 116023, China; (3) College of Animal Technology and University, Northwest Sci.-Tech. University of Agriculture and Forestry, Yangling, Shaanxi 712100, China

This is contribution number G199901209 of 973 from the Chinese basic Chinese BASIC (中文培碁) is the name given to several Chinese-localized versions of the BASIC programming language in the early 1980s. Versions  research project.

* Corresponding author. E-mail: jhxiang@ms.qdio.ac.cn
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Author:Jianhai, Xiang
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