Genetic parameters of growth and survival in the Pacific oyster Crassostrea gigas

The Pacific oyster Crassostrea gigas is a representative bivalve mollusc that is widely cultured in the world and is the largest molluscan group cultured in China. In order to assess the feasibility of improving survival of C. gigas through genetic selection, the heritability and genetic correlations for growth and survival traits between different life stages were examined. Genetic parameters were estimated based on intraclass correlations of 49 full‐sib families (29 half‐sib families) in larvae (4 and 20 days after fertilization) and spat (140 days after fertilization) stages. The heritability for growth traits in larvae and spat was 0.30–0.86 and 0.53–0.59, respectively, and varied with ages. The heritability of survival was low in larvae (0.13 ± 0.05 and 0.17 ± 0.04, respectively for 4 and 20 days after fertilization) but medium (0.39 ± 0.07) in spat, suggesting that selection for increasing spat survival was feasible. The genetic correlation between growth traits within age was medium to high and positive (ranging from 0.47 to 0.96, respectively, between shell length (SL) and shell height (SH) at 20 days and between SL and SH at 140 days after fertilization), suggesting that selection to improve single growth trait will cause positive response in another growth traits in C. gigas. The genetic correlations between survival and growth traits at 140 days were low but positive (ranging from 0.23 to 0.27, respectively, between survival and SH and between survival and SL at 140 days after fertilization), suggesting that selection for survival may not have a negative response in growth. Overall, this study suggests that survival traits should be taken as improving target of next selection breeding programme in C. gigas.     

栉江珧形态性状对重量性状的影响

随机选取101只野生栉江珧,测量其壳长(SL)、壳宽(SW)、壳高(SH)、壳顶角(AG)4个形态性状和体重(WG)、软体重(WT)、闭壳肌重(WM)3个重量性状,采用相关分析和通径分析方法得出了不同形态性状对重量性状的影响效果.结果表明,除壳顶角与闭壳肌重的相关系数不显著外,其余形态性状与重量性状间的相关系数均达极显著水平(P＜0.01);壳高对重量性状的直接作用和间接作用均较大,是影响重量性状的主要因素;壳长和壳宽通过壳长对重量性状的间接作用较大,直接作用较小,是影响重量性状的两个次要因素;利用逐步回归分析建立了栉江珧形态性状关于体重(Y=69.112SH+52.823SW-751.367)、软体重(Y=37.161SH+43.404SW+4.111SA-614.096)和闭壳肌重(Y=4.275SH-18.610)的最优回归方程.     

Heritability estimates for nutritional quality‐related traits of the Pacific oyster,Crassostrea gigas

Pacific oysters, Crassostrea gigas, are the most abundantly harvested shellfish in the world and are ecologically significant. The content of nutrients, including protein, glycogen, lipid, zinc (Zn), and selenium (Se), is important for oyster meat quality, but heritability estimates of such traits have rarely been reported. In this study, 64 full‐sib families were generated using a nested mating design. Finally, 18 full‐sib families, of which there were nine half‐sib families, with each containing 2 full‐sib families, were sampled for heritability estimates. The narrow‐sense heritabilities of glycogen, protein, lipid, Zn, and Se contents were 0.29 ± 0.02, 0.38 ± 0.02, 0.58 ± 0.08, 0.02 ± 0.02, and 0, respectively. Negative genetic correlations existed between both glycogen and protein content (?0.95 ± 0.004) and between lipid and protein content (?0.59 ± 0.05), whereas a positive correlation was observed between lipid and glycogen content (0.16 ± 0.06). Weak genetic and phenotypical correlations (r = 0–0.2) were observed between shell height and nutritional quality traits. These data demonstrated that glycogen, protein, and lipid content can be chosen in a selective breeding program, but glycogen and lipids cannot be selected together with protein. Furthermore, performing indirect selective breeding for quality traits by selecting traits related to growth is impossible. This study provides information for the development of breeding strategies for oyster quality traits.     

华贵栉孔扇贝数量性状的相关性及通径分析

为查明华贵栉孔扇贝（Chlamys nobilis）数量性状间的相关性,随机选取526只扇贝,对其形态性状（壳高、壳长、绞合线长和壳厚）和质量性状（总质量、贝柱鲜质量、鲜肉质量、干肉质量和贝柱干质量）进行测量,采用多元回归与通径分析方法研究了5个易测数量性状（壳高、壳长、绞合线长、壳厚和总质量）对鲜肉质量、贝柱鲜质量和贝柱干质量的影响。结果表明,总质量与鲜肉质量、贝柱鲜质量和叭柱干质量呈极显著的正相关（P〈0．01）。多元回归与通径分析显示,这5个易测数量性状指标中,总质量对3个经济性状的正面影响非常大,其他指标的影响较小甚至不明显。     

合浦珠母贝不同壳色选育系F6数量性状的相关性和通径分析

为深入了解不同壳色合浦珠母贝(Pinctada fucata)形态性状与体重间的关系,为亲贝的选择提供策略,本研究使用合浦珠母贝传统壳色、金壳色、白壳色、红壳色和黑壳色5种壳色选育系F6各1000只个体的性状数据进行通径分析和聚类分析。测量的性状包括壳长(SL,mm)、壳高(SH,mm)、壳宽(SW,mm)、绞合线长(HL,mm)和体重(BW,g)。结果显示,5种壳色合浦珠母贝所有形态性状与体重间的相关系数均达到极显著(P0.01)。通径分析表明,壳长对传统壳色和白壳色贝体重的直接影响最大,壳高对金壳色和红壳色贝体重的直接影响最大,壳宽对黑壳色贝体重的直接影响最大。利用多元回归方法构建不同壳色合浦珠母贝形态性状与体重(BW,g)间的回归方程,其中传统壳色:BW=-24.999+0.467SL+0.387SH-0.259SW+0.244HL;金壳色:BW=-38.661+0.305SL+0.477SH+0.242SW+0.375HL;白壳色:BW=-4.130+0.332SL+0.300SH-0.307SW+0.088HL;红壳色:BW=-27.307+0.327SL+0.321SH+0.252SW+0.305HL;黑壳色:BW=-40.921+0.278SL+0.335SH+1.076SW+0.269HL。聚类分析显示,传统壳色与金壳色形态特征相近,而白壳色、红壳色和黑壳色形态特征相近。     

Heritability estimates for growth‐related traits in the Pacific oyster (Crassostrea gigas) using a molecular pedigree

Pacific oyster is one of the leading species in world aquaculture, but heritability estimation applying mixed‐family approach has not been actively pursued. In this study, heritability for growth‐related traits in the Pacific oyster was first estimated by creating a single cohort of 45 families in a full‐factorial mating design consisting of nine sires and five dams. A total of 270 offspring were analysed and parentage assignment inferred by six microsatellite markers achieved 100% success. All parents contributed to the spawn and a total of 42 full‐sib families were represented. Using an animal model, heritability estimates at 12 months of age were 0.49 ± 0.25 for shell height, 0.36 ± 0.19 for shell length, 0.45 ± 0.23 for shell width and 0.35 ± 0.17 for wet weight. Genetic correlation between shell height and wet weight was quite high (0.79 ± 0.25), suggesting that direct selection of shell height, which is an easily measurable trait, also improves wet weight. The results obtained in this study indicate that growth‐related traits could be improved by exploiting additive genetic effects through selective breeding.     

不同地理群体菲律宾蛤仔表型性状的相关性与通径分析

随机选取繁殖时期的南方群体北方养殖菲律宾蛤仔(SN)、北方土著品种(NN)和南方群体南方养殖蛤仔(SS),测量其壳长(SL)、壳高(SH)、壳宽(SW)、活体质量(mL)、软体质量(mM)和烘干后软体部干质量(mF),采用相关与通径分析方法分析不同地理群体的菲律宾蛤仔表型性状对软体质量的作用效果,为菲律宾蛤仔的选择育种提供理论依据和测度指标。试验结果表明,不同地理群体菲律宾蛤仔的形态学指标无显著差异(P>0.05);不同地理群体各数量性状之间的相关关系均达到极显著水平(P<0.01);决定系数分析表明,不同地理群体菲律宾蛤仔影响软体质量的主要表型性状并不相同,南方群体北方养殖蛤仔群体的壳宽直接作用最大,壳宽通过壳高的间接作用是次要因素;北方土著品种群体壳长是主要因素,壳长通过壳高的间接作用是次要因素;南方群体南方养殖蛤仔群体壳高的直接作用是主要因素,壳长通过壳高的间接作用是次要因素。对通径系数检验不显著的自变量进行删除,利用逐步回归的方法,建立不同地理群体菲律宾蛤仔软体质量的最优回归方程:SN群体,mM1=-4.276+0.211SH+0.116SW;NN群体,mM2=-2.806+0.088SL+0.087SH;SS群体,mM3=-3.101+0.290SH。回归方程分析表明回归关系均达到极显著水平(P<0.01)。     

Heritability estimates of larval and spat shell height in the Chilean blue mussel (Mytilus chilensis Hupe 1854) produced under controlled laboratory conditions

The heritability of larval size is estimated in Mytilus chilensis, based on a nested design, involving 95 full‐sib and 19 half‐sib families (19 males each mated to 5 separated females) grown under controlled laboratory conditions. The half‐sib heritability estimates at 10, 25 and 40 days for larval and spat shell height ranged between 0.38±0.33 and 0.84±0.45, suggesting that selection for shell size would be effective.     

西施舌形态性状与体质量性状的相关性分析

为研究西施舌(Coelomactra antiquata)野生群体壳体性状对体质量性状的影响,文章以广西北海铁山港海区采集的178只西施舌为研究对象,测量壳高(SH)、壳长(SL)、壳宽(SW)、前缘长(AL)、后缘长(GL)、壳质量(SM)、软体部质量(VM)、活体质量(BM),以壳体性状(SH、SL、SW、AL、GL、SM)为自变量,体质量性状(BM、VM)为因变量进行相关性分析、通径分析、多元回归分析。结果表明西施舌8个数量性状中,软体部质量变异系数最高,性状间的相关性均达到极显著水平(P<0.01),活体质量和软体部质量分别与壳高和壳质量的相关系数最大(0.831和0.646);壳宽和壳质量分别对活体质量和软体部质量的直接影响最大,通径系数分别为0.362和0.487,决定程度分析与通径分析结果变化趋势一致;建立了关于体质量性状的回归方程(R2<0.85)。     

扇贝新品系“紫海墨”F4的生长特征和生产性能分析

为考察扇贝新品系"紫海墨"的生产性能,为新品系选育效果提供依据,在同一海域养殖、相同管理模式和相同养殖周期下,对新品系"紫海墨"F4与墨西哥湾扇贝(Argopecten irradians concentricus)养殖群体的壳长、壳高、壳宽、体质量、闭壳肌质量和壳质量等7个性状及各阶段成活率进行生长特征和生产性能分析。结果表明,新品系"紫海墨"F4从10月13日养殖至翌年4月28日,壳长从1~1.5 mm长至50~60 mm的成品时,累积成活率为51.32%,显著大于墨西哥湾扇贝的29.18%(P<0.01);养殖周期内,两个养殖群体同一时期的壳长、壳高、壳宽和体质量也有极显著差异(P<0.01),新品系"紫海墨"F₄的生长速度高于墨西哥湾扇贝;经过6.5个月的养殖,新品系"紫海墨"F₄的壳长、壳高、壳宽、体质量和闭壳肌质量分别比墨西哥湾扇贝提高了19.50%、19.84%、16.23%、56.30%和20.20%。     

Heritability of larval and juvenile growth for two stocks of Manila clam Ruditapes philippinarum

The heritability of larval and juvenile growth in natural and cultured populations of the Manila clam Ruditapes philippinarum was estimated using an unbalanced nested design and an artificial fertilization technique. A total of 33 full‐sib families and 10 half‐sib families were obtained from the natural population in Shihe (Dalian, Liaoning Province in the north‐eastern part of China), whereas 11 half‐sib families and 33 full‐sib families were generated from the cultured population in Putian (Fujian Province in the south‐eastern part of China). The offspring from each family were reared under the same hatchery and nursery conditions. Results from these studies led to estimates indicating a significantly greater influence due to the dam component compared with the sire component for both natural and cultured populations, which was mainly due to maternal effect. We found that the heritability estimates obtained from the intra‐group correlation of sire half‐sib were precise and unbiased. The narrow‐sense heritabilities of shell length for the Manila clam in the natural population (NP) at larval (at the age of 9 days) and juvenile (at the age of 30 days) stages were 0.22 ± 0.11 and 0.39 ± 0.14, respectively, whereas those in the cultured population (CP) were 0.17 ± 0.09 and 0.87 ± 0.24 respectively. The heritabilities of both NP and CP at 3 days were not significantly different from zero, and were ?0.03 for NP and ?0.14 for CP. These results suggest that selection should be highly effective in Manila clam, and that selecting either a natural or a cultured population for producing faster growing larvae and juveniles should be successful. Selective breeding could thus be used to develop high‐quality seed that would facilitate the development of a shorter culture cycle and higher yields.     

Growth of wild pearl oysters <Emphasis Type="Italic">Pinctada fucata,Pinctada margaritifera</Emphasis> and <Emphasis Type="Italic">Pinctada sugillata</Emphasis> (Bivalvia: Pteriidae) in Taiwan

In order to understand growth features of pearl oysters in the genus Pinctada, i.e. Pinctada fucata, Pinctada margaritifera, and Pinctada sugillata in Taiwan, a total of 3062 wild individuals of these species from juvenile to adult were collected monthyly from March 2001 to April 2002 in Jukeng, Pingtung County, south-west Taiwan. Quantitative measurements of live oysters were conducted for shell height (SH), shell length (SL), shell width (SW), hinge length (HL), and wet weight (WW). Different cohorts were identified through multiple length frequency analysis on SH of P. fucata and P. margaritifera, and growth curves with seasonal variation were estimated for these species. Pinctada fucata in Taiwan had a different seasonal growth pattern from the Japanese population, but had similar growth rates during the high growth period. The growth rate of P. margaritifera in Taiwan was slower than in French Polynesia, the Solomon Islands, and the Red Sea. Comparisons of morphological growth features among the three species show large differences in the SW-related features. Pinctada fucata in Taiwan had larger SW than in Japan and Korea. The differences in growth rates and morphological features suggested that the wild Taiwanese oysters may retain genetically pristine characteristics, thus genetic conservation might be urgently needed.     

Family growth response to different laboratory culture environments shows genotype–environment interaction in the sea urchin Strongylocentrotus intermedius

This study investigated the genetic variation and genotype–environment interaction (GEI) in the early growth stage among seven full‐sib families of the sea urchin Strongylocentrotus intermedius. From each of the seven families, 180 or 270 sea urchins of the same size were exposed to three laboratory environments (E₁, E₂ and E₃). These environments were commonly used in breeding programs for S. intermedius. After a 102 days trial, test height, test diameter, body weight and the coefficient of variation of body weight were determined. The results revealed significant family effects on most growth traits in all the three environments (except for test diameter in environment E₂), and significant environmental effects on growth in several families (family 1, 5, 6, 7). The study also revealed that the coefficient of variation in body weight varied significantly among the families (P < 0.01) but not among the environments. Highly significant GEI effects were also recorded for all growth traits (P ≤ 0.001), except for the coefficient of variation in body weight (P > 0.05). Variances of GEI accounted for about 0–2.742% of phenotypic variances for the investigated traits. Significant GEI caused a certain degree family ranking inversions for the growth traits. The present study provides evidence for the existence of GEI in the family selection of sea urchin S. intermedius. More attention should be paid to the GEI to obtain satisfactory genetic gain in S. intermedius.     

Breeding for improved resistance to Edwardsiella ictaluri in striped catfish (Pangasianodon hypophthalmus): Quantitative genetic parameters

Bacillary necrosis of Pangasius (BNP), caused by Edwardsiella ictaluri, is one of the most devastating diseases in striped catfish farming. To date, quantitative genetic inheritance of BNP resistance is not known in striped catfish Pangasianodon hypophthalmus. The main aim of this study was to estimate genetic parameters for BNP resistance in a breeding population of striped catfish undergoing four generations of selection for high growth. Specifically, the study examined whether BNP resistance is heritable to enable family selection and whether genetic improvement for enhanced BNP resistance may have detrimental effects on growth and survival rate. To test these hypotheses, 720 full‐ and half‐sib families were challenged with E. ictaluri pathogen using injection and cohabitation methods over four years, from 2010 to 2012 and 2015. In total, the data included 398,234 animals in the pedigree, from which 18,849 animals had disease challenge test records and 39,103 siblings had growth performance. Both univariate and bivariate sire–dam linear and threshold mixed models were used to estimate (co)variance components for BNP resistance, survivals and growth traits. The estimates of heritability for the BNP resistance recorded as death or survival were low regardless of models used (0.10–0.16), whereas survival time (days post‐challenge test) showed moderate heritability (0.35). The survival rate during hapa rearing had medium heritability (0.33–0.52). The genetic correlations of BNP resistance with body weight and survival were all positive (0.03–0.53), suggesting that selection of increased BNP resistance may have positive impacts on growth and survival traits, and these traits could be easily improved simultaneously in the selective breeding programme for striped catfish.     

鳙30日龄生长性状的遗传参数

为了研究鳙早期生长性状的遗传改良潜力,利用2个亲本群体进行群体繁殖(组1)和人工授精(组2)实验,并分别采集672尾30日龄鱼苗,用于开展对早期生长性状的遗传分析。通过微卫星标记分别鉴定了其中628和660尾个体的亲本来源,并依此获取群体双列杂交的信息;亲本对应子代贡献率存在极显著差异。在2个繁殖组中,体质量和体长在家系间均存在极显著差异;在组2中,体质量和体长在交配设计间均存在显著差异。此外,杂交组合的2个生长性状均呈现出中亲杂种优势(0.39%~7.64%),其特殊配合力也均为正值(0.01~0.02)。基于动物模型和限制性最大似然法,鳙30日龄体质量和体长的遗传力估值分别为0.47和0.49,且均达到极显著水平。体质量和体长间存在极显著的遗传相关和表型相关,分别为0.89和0.83。研究表明,通过家系构建和群体杂交,可以获得具有生长优势的鳙鱼苗;鳙30日龄生长性状具有较高选育潜力,而亲本对子代贡献的不平衡现象在选育过程中需要引起重视。     

Quantitative genetic properties of four measures of deformity in yellowtail kingfish Seriola lalandi Valenciennes, 1833

The main aim of this study was to estimate the heritability for four measures of deformity and their genetic associations with growth (body weight and length), carcass (fillet weight and yield) and flesh‐quality (fillet fat content) traits in yellowtail kingfish Seriola lalandi. The observed major deformities included lower jaw, nasal erosion, deformed operculum and skinny fish on 480 individuals from 22 families at Clean Seas Tuna Ltd. They were typically recorded as binary traits (presence or absence) and were analysed separately by both threshold generalized models and standard animal mixed models. Consistency of the models was evaluated by calculating simple Pearson correlation of breeding values of full‐sib families for jaw deformity. Genetic and phenotypic correlations among traits were estimated using a multitrait linear mixed model in ASReml. Both threshold and linear mixed model analysis showed that there is additive genetic variation in the four measures of deformity, with the estimates of heritability obtained from the former (threshold) models on liability scale ranging from 0.14 to 0.66 (SE 0.32–0.56) and from the latter (linear animal and sire) models on original (observed) scale, 0.01–0.23 (SE 0.03–0.16). When the estimates on the underlying liability were transformed to the observed scale (0, 1), they were generally consistent between threshold and linear mixed models. Phenotypic correlations among deformity traits were weak (close to zero). The genetic correlations among deformity traits were not significantly different from zero. Body weight and fillet carcass showed significant positive genetic correlations with jaw deformity (0.75 and 0.95, respectively). Genetic correlation between body weight and operculum was negative (?0.51, P < 0.05). The genetic correlations' estimates of body and carcass traits with other deformity were not significant due to their relatively high standard errors. Our results showed that there are prospects for genetic selection to improve deformity in yellowtail kingfish and that measures of deformity should be included in the recording scheme, breeding objectives and selection index in practical selective breeding programmes due to the antagonistic genetic correlations of deformed jaws with body and carcass performance.     

虾夷扇贝獐子岛红品系家系建立及生长发育的比较

采用不平衡巢式设计(♂∶♀=1∶3),建立了12个金贝广场挑选的獐子岛红虾夷扇贝父系半同胞(A、B、C……L)和36个母系全同胞(A1、A2、A3,B1、B2、B3,C1、C2、C3……L1、L2、L3)家系,比较各家系间浮游幼虫期和稚贝期的生长及存活指标,以评估虾夷扇贝"獐子岛红"品系的生长性状。结果表明,K3家系5日龄幼虫生长速度最快,比生长最慢的B3平均壳长大9.40%;I1家系在15日龄时生长速度最快,壳长为164.83μm,C3、D3家系25日龄壳长最大,较平均壳长高17.71%;D2家系120日龄稚贝壳高生长最快,较生长最慢的C1和D1家系平均壳高大40.41%;D2家系180日龄壳高依然最高,较H1平均壳高(18.3mm)高38.96%。I1家系幼虫存活率最高,较D1家系平均存活率高17.67%;F2家系稚贝存活率最高,显著高于其他试验组(P0.05,n=108)。综合36个同胞家系的各指标,D2、I1家系生长、抗逆性状最优,可作为优良品种选育的育种材料。     

Genetic parameters for growth‐related traits and survival with age in the Kuruma shrimp Marsupenaeus japonicus

Genetic parameters were estimated for growth‐related traits and survival of the Kuruma shrimp Marsupenaeus japonicus based on 66 families, including 30 paternal half‐sib families, which were obtained using artificial insemination of two females by each male. The variance components for growth‐related trait and survival were estimated using a single‐trait animal model and a sir‐dam model, respectively, and genetic parameters were estimated using the restricted maximum likelihood method. The coefficients of variation for growth‐related traits such as body length (BL), abdominal length (AL), and body weight (BW) were 9.36%–22.44%, 10.61%–21.92%, and 26.47%–58.33%, respectively, at different ages (45, 75, 105, 135, and 165 days). The corresponding heritability estimates for each growth trait were 0.1545–0.1951, 0.1672–0.1905, and 0.1596–0.1934, respectively, all of which were found to be at moderate levels and increased with age. The heritability of survival on day 165 was low (0.003). The genetic correlations among growth traits were positive and high (0.7316–0.9896) at the different ages, suggesting that selection to improve any single growth trait will cause positive responses in other growth traits examined in M. japonicus. The genetic correlations between growth traits and survival were also positive but low (0.005–0.087), which indicated that only selecting for growth traits may not cause a positive correlated response in the survival of the core population. According to the above results, we suggested that growth and survival traits should be taken as improving targets of breeding in M. japonicus. These results provided reference data for selective breeding and multitrait selection of M. japonicus.     

长牡蛎生长性状遗传力、遗传相关和表型相关分析

为了查清乳山(RS)和崆峒岛(KTD)海域长牡蛎生长性状各指标的遗传力、遗传相关和表型相关,本实验通过部分因子交配设计和人工授精的方法建立家系,并在乳山和崆峒岛海域进行养殖。用混合线性模型分别计算RS和KTD海域188日龄、338日龄和474日龄长牡蛎生长性状各指标的遗传参数。结果发现:不同日龄长牡蛎生长性状各指标,在乳山海域的遗传力是0.28~0.55,为中高遗传力,在崆峒岛海域的遗传力是0.34~0.63,为高遗传力;与乳山海域相比,崆峒岛海域各指标的遗传力偏高。运用亲本模型,将日龄和地点作为固定效应,计算得到壳高、壳长、壳宽和湿重的遗传力分别为0.25±0.08、0.29±0.09、0.14±0.05和0.26±0.09。不同海域、不同日龄的遗传相关和表型相关结果各不相同,但各指标间均呈现正相关;总体来讲,性状间的遗传相关大于表型相关。实验结果有助于掌握山东半岛南北两侧长牡蛎生长性状的数量遗传学参数,为制定长牡蛎在该海域的选育技术路线提供数据支持。     

Genetic variation in the growth traits of straight-bred and crossbred black bream (Acanthopagrus butcheri Munro) at 90 days of age

The current slow growth rate of black bream (Acanthopagrus butcheri Munro) impedes their widespread commercial aquaculture across inland southern Australia. We report initial estimates of genetic variation for growth traits at 90 days of age from a diallel cross using two black bream populations. Standard length, total length, and wet weight varied significantly among lines and among half‐sib groups within lines. Differences among half‐sib groups explained 6.8% of the total variance in standard length, 8.3% in total length, and 7.1% in wet weight, giving estimated heritabilities over all lines of 0.27±0.11 for standard length, 0.33±0.13 for total length, and 0.28±0.12 for wet weight. There was no evidence for heterosis in any traits when straight‐bred and crossbred lines were compared. There were high phenotypic (r_P=0.95–0.98) and genetic (r_G=0.63–0.69) correlations among all growth traits.