Explaining bathymetric diversity patterns in marine benthic invertebrates and demersal fishes: physiological contributions to adaptation of life at depth.

Explaining bathymetric diversity patterns in marine benthic invertebrates and demersal fishes: physiological contributions to adaptation of life at depth.
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DOI:
10.1111/brv.12061
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发表时间:
2014-05
影响因子:
--
通讯作者:
Thatje S
Thatje S
中科院分区:
其他
文献类型:
--
作者:
Brown A;Thatje S

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在深大陆边缘现存的动物群中,确定了海洋底栖无脊椎动物和底栖鱼类的深海生物多样性格局。从陆架断裂到1000 m,陆架动物群和斜坡动物群之间的过渡,以及从2000到3000 m,斜坡动物群和深海动物群之间的过渡,是广泛而明显的深度分带;这些转变的特点是高物种周转率。多样性呈单峰模式,深度在1000 ~ 3000 m之间,尽管这些深度所代表的面积相对较小。分带被认为是在显生宙的多次大灭绝事件之后,浅水生物在深海定居的结果。低温和高压的影响跨越了生物组织的等级水平,似乎足以限制这些浅水物种的分布。在实验中,深海的静水压力一直被确定为浅水和深海底栖无脊椎动物在原位温度下所能承受的最大压力,而底栖无脊椎动物和底栖鱼类似乎都需要适应这种压力才能进入更深的水域。总之,这表明深海深处的高压和热生理瓶颈有助于深海分带。单峰多样性-深度模式的峰值通常出现在这些深度,尽管这些深度所代表的面积相对较低。虽然人们认识到,在漫长的进化时间尺度上,浅水多样性模式是由物种形成驱动的,但很少考虑到物种分布模式随深度的潜在影响。分子和形态学证据表明,凉爽的深海水域是深海适应性辐射的主要场所,我们假设物种形成率的水深变化可能随着时间的推移驱动单峰多样性-深度模式。对代谢率依赖性突变和代数的热效应已被提出驱动物种形成率的差异,这导致了现代纬度生物多样性模式随时间的变化。显然,这种热机制本身不能解释水深模式,因为温度通常随深度而降低。我们假设,深海深处的高静水压力和低温对入侵深海的浅水类群的生理影响,可能通过增加生殖细胞的诱变活性,在胚胎或幼虫发育期间灭活管道,释放隐藏的变异或诱变活性,或通过激活或释放幼虫或成虫的转座因子,调用应激进化机制。在这种情况下,深海深处生理瓶颈的变异增加导致物种形成率升高。提高对高静水压力和低温的耐受性的适应,使深海深处的定植和减少应力-进化反应,从而使深海分类群的物种形成恢复到背景速率。随着时间的推移,这种机制可能导致单模态多样性-深度模式。
Bathymetric biodiversity patterns of marine benthic invertebrates and demersal fishes have been identified in the extant fauna of the deep continental margins. Depth zonation is widespread and evident through a transition between shelf and slope fauna from the shelf break to 1000 m, and a transition between slope and abyssal fauna from 2000 to 3000 m; these transitions are characterised by high species turnover. A unimodal pattern of diversity with depth peaks between 1000 and 3000 m, despite the relatively low area represented by these depths. Zonation is thought to result from the colonisation of the deep sea by shallow-water organisms following multiple mass extinction events throughout the Phanerozoic. The effects of low temperature and high pressure act across hierarchical levels of biological organisation and appear sufficient to limit the distributions of such shallow-water species. Hydrostatic pressures of bathyal depths have consistently been identified experimentally as the maximum tolerated by shallow-water and upper bathyal benthic invertebrates at in situ temperatures, and adaptation appears required for passage to deeper water in both benthic invertebrates and demersal fishes. Together, this suggests that a hyperbaric and thermal physiological bottleneck at bathyal depths contributes to bathymetric zonation. The peak of the unimodal diversity–depth pattern typically occurs at these depths even though the area represented by these depths is relatively low. Although it is recognised that, over long evolutionary time scales, shallow-water diversity patterns are driven by speciation, little consideration has been given to the potential implications for species distribution patterns with depth. Molecular and morphological evidence indicates that cool bathyal waters are the primary site of adaptive radiation in the deep sea, and we hypothesise that bathymetric variation in speciation rates could drive the unimodal diversity–depth pattern over time. Thermal effects on metabolic-rate-dependent mutation and on generation times have been proposed to drive differences in speciation rates, which result in modern latitudinal biodiversity patterns over time. Clearly, this thermal mechanism alone cannot explain bathymetric patterns since temperature generally decreases with depth. We hypothesise that demonstrated physiological effects of high hydrostatic pressure and low temperature at bathyal depths, acting on shallow-water taxa invading the deep sea, may invoke a stress–evolution mechanism by increasing mutagenic activity in germ cells, by inactivating canalisation during embryonic or larval development, by releasing hidden variation or mutagenic activity, or by activating or releasing transposable elements in larvae or adults. In this scenario, increased variation at a physiological bottleneck at bathyal depths results in elevated speciation rate. Adaptation that increases tolerance to high hydrostatic pressure and low temperature allows colonisation of abyssal depths and reduces the stress–evolution response, consequently returning speciation of deeper taxa to the background rate. Over time this mechanism could contribute to the unimodal diversity–depth pattern.
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