Ayre D J, Hughes T P
Australian Flora and Fauna Research Centre, Department of Biological Sciences, University of Wollongong, New South Wales.
Evolution. 2000 Oct;54(5):1590-605. doi: 10.1111/j.0014-3820.2000.tb00704.x.
Marine organisms exhibit great variation in reproductive modes, larval types, and other life-history traits that may have major evolutionary consequences. We measured local and regional patterns of genetic variation in corals along Australia's Great Barrier Reef to determine the relative contributions of sexual and asexual reproduction to recruitment and to infer levels of gene flow both locally (among adjacent sites, < 5 km apart) and regionally (among reefs separated by 500-1,200 km). We selected five common brooding species (Acropora cuneata, A. palifera, Pocillopora damicornis, Seriatopora hystrix, and Stylophora pistillata) and four broadcast spawners (Acropora hyacinthus, A. cytherea, A. millepora, and A. valida), which encompassed a wide range of larval types and potential dispersal capabilities. We found substantial genotypic diversity at local scales in six of the nine species (four brooders, two spawners). For these six, each local population displayed approximately the levels of multilocus genotypic diversity (Go) expected for outcrossed sexual reproduction (mean values of Go:Ge ranged from 0.85 to 1.02), although consistent single-locus heterozygous deficits indicate that inbreeding occurs at the scale of whole reefs. The remaining three species, the brooder S. hystrix and the spawners A. valida and A. millepora displayed significantly less multilocus genotypic diversity (Go) than was expected for outcrossed sexual reproduction (Ge) within each of several sites. Acropora valida and A. millepora showed evidence of extensive localized asexual replication: (1) a small number of multilocus (clonal) genotypes were numerically dominant within some sites (Go:Ge values were as low as 0.17 and 0.20): (2) single-locus genotype frequencies were characterized by both excesses and deficits of heterozygotes (cf. Hardy-Weinberg expectations), and (3) significant linkage disequilibria occurred. For the brooding S. hystrix Go:Ge values were also low within each of four sites (x = 0.48). However, this result most likely reflects the highly restricted dispersal of gametes or larvae, because levels of genetic variation among sites within reefs were extremely high (FSR = 0.28). For all species, we detected considerable genetic subdivision among sites within each reef (high FSR-values), and we infer that larval dispersal is surprisingly limited (i.e., Nem among sites ranging from 0.6 to 3.3 migrants per generation), even in species that have relatively long planktonic durations. Nevertheless, our estimates of allelic variation among reefs (FRT) also imply that for all four broadcast spawning species and three of the brooders, larval dispersal is sufficient to maintain moderate to high levels of gene flow along the entire Great Barrier Reef (i.e., Nem among reefs ranged from 5 to 31). In contrast, widespread populations of S. hystrix and S. pistilata (the two remaining brooders) are relatively weakly connected (Nem among reefs was 1.4 and 2.5, respectively). We conclude that most recruitment by corals is very local, particularly in brooders, but that enough propagules are widely dispersed to ensure that both broadcast spawning and brooding species form vast effectively panmictic populations on the Great Barrier Reef.
海洋生物在繁殖方式、幼体类型及其他生活史特征方面表现出巨大差异,这些差异可能具有重大的进化意义。我们测量了澳大利亚大堡礁珊瑚的局部和区域遗传变异模式,以确定有性繁殖和无性繁殖对补充的相对贡献,并推断局部(相邻地点之间,相距小于5公里)和区域(相距500 - 1200公里的珊瑚礁之间)的基因流水平。我们选择了5种常见的产卵珊瑚(楔形鹿角珊瑚、帕氏鹿角珊瑚、鹿角杯形珊瑚、刺星珊瑚和扁脑珊瑚)和4种排卵珊瑚(风信子鹿角珊瑚、赛氏鹿角珊瑚、多孔鹿角珊瑚和强壮鹿角珊瑚),它们涵盖了广泛的幼体类型和潜在扩散能力。我们发现9种珊瑚中有6种(4种产卵珊瑚、2种排卵珊瑚)在局部尺度上具有丰富的基因型多样性。对于这6种珊瑚,每个局部种群显示出的多位点基因型多样性(Go)水平与异交有性繁殖预期的水平相近(Go:Ge的平均值在0.85至1.02之间),尽管单一基因座杂合子的持续不足表明在整个珊瑚礁尺度上存在近亲繁殖。其余3种珊瑚,产卵的刺星珊瑚以及排卵的强壮鹿角珊瑚和多孔鹿角珊瑚,在几个地点中的每个地点,其多位点基因型多样性(Go)均显著低于异交有性繁殖预期的水平(Ge)。强壮鹿角珊瑚和多孔鹿角珊瑚显示出广泛的局部无性复制证据:(1)少数多位点(克隆)基因型在某些地点数量上占主导(Go:Ge值低至0.17和0.20);(2)单一基因座基因型频率的特征是杂合子过剩和不足(与哈迪 - 温伯格预期相比),并且(3)出现了显著的连锁不平衡。对于产卵的刺星珊瑚,在4个地点中的每个地点Go:Ge值也很低(x = 0.48)。然而,这一结果很可能反映了配子或幼体的扩散受到高度限制,因为珊瑚礁内各地点之间的遗传变异水平极高(FSR = 0.28)。对于所有物种,我们在每个珊瑚礁内的不同地点检测到了相当程度的遗传分化(高FSR值),并且我们推断幼体扩散令人惊讶地有限(即各地点之间每代的迁移数Nem在0.6至3.3之间),即使在浮游期相对较长的物种中也是如此。尽管如此,我们对珊瑚礁间等位基因变异的估计(FRT)也表明,对于所有4种排卵珊瑚物种和3种产卵珊瑚中的3种,幼体扩散足以在整个大堡礁维持中等至高的基因流水平(即珊瑚礁间的Nem在5至31之间)。相比之下,广泛分布的刺星珊瑚和扁脑珊瑚种群(剩下的2种产卵珊瑚)之间的联系相对较弱(珊瑚礁间的Nem分别为1.4和2.5)。我们得出结论,珊瑚的大多数补充是非常局部性的,特别是在产卵珊瑚中,但有足够的繁殖体广泛扩散,以确保排卵珊瑚和产卵珊瑚物种在大堡礁上都能形成庞大的有效随机交配种群。