Levin Lisa A, Zhu Jun, Creed Elizabeth
Department of Marine, Earth and Atmospheric Sciences, Raleigh, NC, 27695-8208, USA.
Department of Statistics, North Carolina State University, Raleigh, NC, 27695-8208, USA.
Evolution. 1991 Mar;45(2):380-397. doi: 10.1111/j.1558-5646.1991.tb04412.x.
The polychaete Streblospio benedicti is unusual in that several field populations consist of individuals that exhibit either planktotrophic or lecithotrophic larval development. Planktotrophy in this species involves production of many small ova that develop into feeding larvae with a two- to three-week planktonic period. Lecithotrophy involves production of fewer, larger ova that develop into nonfeeding larvae that are brooded longer and have a brief planktonic stage. Reciprocal matings were performed to investigate genetic variance components and the correlation structure of life-history traits associated with planktotrophy and lecithotrophy. Our objective was to better understand persistence of this developmental dichotomy in Streblospio benedicti, and among marine invertebrates in general. Substantial additive genetic variation (75-98% of total) was detected for the following characters at first reproduction: female length; position of the first gametogenic setiger and first brood pouch; ovum diameter; three traits related to fecundity (numbers of ova per ovary, larvae per brood pouch, and larvae per brood); median larval planktonic period and the presence of larval swimming setae; but not for total number of brood pouches; larval length; larval feeding; and larval survivorship. Based on the unusual geographic distribution of development modes in this species, we hypothesize that the developmental traits have evolved in allopatry and have only recently come into contact in North Carolina. The high additive contribution to variance observed for many traits may be inflated due to (a) nonrandom breeding in nature, and (b) examination of only one component of an age-structured population at one time. Nuclear interaction variance and maternal variance accounted for 84% of the total variation in larval survivorship. This observation supports other empirical studies and theoretical predictions that nonadditive components of variance will increase in importance in individual traits that are most closely tied to fitness. A network of life-history trait correlations was observed that defines distinct planktotrophic and lecithotrophic trait complexes. Negative genetic correlations were present between fecundity and egg size, between fecundity and position of the first gametes, and between larval survivorship and median planktonic period. Positive genetic correlations were detected between fecundity and female size at first reproduction and between planktonic period and the presence of swimming setae. Intergenerational product-moment correlations were negative for larval length and fecundity, planktonic period and egg size, female size and larval survivorship, and fecundity and larval survivorship. If the genetic correlation structure observed in the laboratory persists in the field, it may constrain responses of individual characters to directional selection, and indirectly perpetuate the dichotomies associated with planktotrophy and lecithotrophy.
多毛纲动物贝氏斯特雷螺(Streblospio benedicti)很不寻常,因为在几个野外种群中,个体呈现出不同的发育方式,要么是浮游生物营养型幼体发育,要么是卵黄营养型幼体发育。该物种的浮游生物营养型发育方式涉及产生许多小卵,这些小卵发育成摄食性幼体,有两到三周的浮游期。卵黄营养型发育方式则是产生较少、较大的卵,这些卵发育成非摄食性幼体,幼体在亲体处停留时间更长,且有短暂的浮游阶段。我们进行了正反交实验,以研究与浮游生物营养型和卵黄营养型发育相关的遗传方差成分以及生活史性状的相关结构。我们的目标是更好地理解贝氏斯特雷螺以及一般海洋无脊椎动物中这种发育二分法的持续性。在首次繁殖时,检测到以下性状存在大量加性遗传变异(占总变异的75 - 98%):雌体长度;首个配子发生刚节和首个育卵囊的位置;卵直径;与繁殖力相关的三个性状(每个卵巢的卵数、每个育卵囊的幼体数以及每次育幼的幼体数);幼体浮游期的中位数以及幼体游泳刚毛的有无;但未检测到育卵囊总数、幼体长度、幼体摄食以及幼体存活率的加性遗传变异。基于该物种发育模式不寻常的地理分布,我们推测这些发育性状是在异域进化的,并且直到最近才在北卡罗来纳州接触到。许多性状观察到的高加性方差贡献可能由于以下原因而被夸大:(a)自然界中的非随机交配,以及(b)一次仅检查年龄结构种群的一个组成部分。核相互作用方差和母体方差占幼体存活率总变异的84%。这一观察结果支持了其他实证研究和理论预测,即方差的非加性成分在与适合度最密切相关的个体性状中重要性将增加。观察到一个生活史性状相关网络,该网络定义了不同的浮游生物营养型和卵黄营养型性状复合体。繁殖力与卵大小之间、繁殖力与首个配子位置之间以及幼体存活率与浮游期中位数之间存在负遗传相关性。在首次繁殖时,繁殖力与雌体大小之间以及浮游期与游泳刚毛的有无之间检测到正遗传相关性。幼体长度与繁殖力、浮游期与卵大小、雌体大小与幼体存活率以及繁殖力与幼体存活率之间的代际积矩相关性为负。如果在实验室中观察到的遗传相关结构在野外持续存在,它可能会限制个体性状对定向选择的响应,并间接使与浮游生物营养型和卵黄营养型相关的二分法永久化。