生物多样性 ›› 2020, Vol. 28 ›› Issue (7): 821-832. DOI: 10.17520/biods.2019312
收稿日期:
2019-10-22
接受日期:
2020-02-22
出版日期:
2020-07-20
发布日期:
2020-09-29
通讯作者:
杨锡福
作者简介:
* E-mail: yangxifu@ioz.ac.cn基金资助:
Xifu Yang1,*(), Hongmao Zhang2, Zhibin Zhang1,3
Received:
2019-10-22
Accepted:
2020-02-22
Online:
2020-07-20
Published:
2020-09-29
Contact:
Xifu Yang
摘要:
大年结实(mast seeding)是多生年植物种群周期性同步大量繁殖的一种自然现象。大年结实作为植物适应环境条件、提高繁殖能力的一种策略而备受关注, 但其驱动机制和进化意义尚存在较大争议。在依赖动物扩散种子的植物中, 大年结实被认为是一种调控动物贮食行为、提高种子扩散效率, 并最终增加繁殖成功率的一种策略; 动物介导的植物间互作可能是促进植物共存的进化驱动力。本文简要梳理了大年结实现象的各种假说, 提出了一个包括气候、资源、动植物互作的理解大年结实机制的概念框架, 并着重讨论了大年结实和动物贮食行为之间的关系及其进化和生态意义。建议未来研究需要借助长期生态监测和分子生物学方法, 揭示植物大年结实与动物贮食行为之间的生态与进化过程。
杨锡福, 张洪茂, 张知彬 (2020) 植物大年结实及其与动物贮食行为之间的关系. 生物多样性, 28, 821-832. DOI: 10.17520/biods.2019312.
Xifu Yang, Hongmao Zhang, Zhibin Zhang (2020) Mast seeding and its relationship to animal hoarding behaviour. Biodiversity Science, 28, 821-832. DOI: 10.17520/biods.2019312.
假说 Hypothesis | 描述 Description | 典型特征 Typical characteristics | 评论 Comment | |
---|---|---|---|---|
资源匹配(气候跟踪) Resource matching (Weather tracking) | 种子产量的变化是由于植物可利用资源的变化造成的 Seed crops vary in response to plant available resources variation ( | 在不改变营养生长或储备资源的情况下实现种子产量的变化 Variable seed crops achieved without switching resources from vegetative growth or reserves | 较早被提出, 早期采用反证法; 多数支持而少数不完全支持 Put forward earlier, and early adopt proof by contradiction ( | |
捕食者饱和 Predator satiation | 大量结实使捕食者饱和, 导致个体被捕食的比例降低 Large seed crops satiate seed predators, which thus lead to individuals experience lower proportion seed predation ( | 种子大年捕食率较低 Lower predation rate in high-seed years | 被多数研究支持; 不过, 取决于功能群的类型 Well supported by many studies ( | |
风媒授粉 Wind pollination | 风媒授粉植物同步大量开花可以获得更高的传粉效率 Wind-pollinated plants can achieve greater pollination efficiency through synchronized above-average flowering effort ( | 在大量开花年受精比例更高 Higher % fertilization in high flowering years | 很好地被验证; 然而, 部分实验结果是模棱两可的 Well documented ( | |
动物扩散(捕食者扩散) Animal dispersal (Predator dispersal) | 大年结实吸引更多扩散者, 因而刺激分散贮藏动物更多的扩散和贮藏种子 Large fruit production attract more seed dispersers, and result in wider dispersal and hoard by scater-hoarders ( | 种子大年扩散率更高或扩散更远 Higher seed dispersal rate or greater dispersal distances in high-seed years | 是否有利于持续繁殖取决于扩散者的功能反应; 部分结论支持, 部分反对 Whether favor constant reproduction, depending on functional response of dispersers; some evidences in support ( | |
环境预测 Environmental prediction | 植物可以“预知”最适气候条件而繁殖和建成 Plants can “predict” favorable climate conditions for reproduction and establishment ( | 种子或幼苗在大年结实时生存率更高 Higher seed or seedling survival in mast seeding | 部分证据支持, 部分反对 Some evidences in support ( | |
大种子 Large seed | 生产大种子增加了种子大年间的恢复时间 Selection for larger seed size increases the recovery time between large seed crops ( | 在种子较大的物种中, 种子数随年际变化更大 More variable seed number among years in related species with large seeds | 部分结论支持, 部分反对; 其他研究认为种子大小和种子数量存在权衡; 中等大小的种子具有更高的建苗率 Some support ( | |
动物传粉 Animal pollination | 大量开花更易于吸引动物传粉者, 从而提高传粉效率 Large flowering attract animal pollinators, thus enhance pollination efficiency ( | 在大量开花年受精比例更高 Higher % fertilization in high-flowering years | 一些分布稀少的植物个体同时产生大量的花, 更有助于种群数量的增加 A few sparsely distributed individuals that simultaneously produce a large number of flowers are more conducive to population growth ( | |
辅助成本 Accessory cost | 较高的固定繁殖成本有利于更少、更大规模的繁殖活动 Higher fixed reproductive costs favor fewer, larger reproductive episodes ( | 在植物内进行大量繁殖时, 每粒种子的辅助成本较低 Lower accessory costs per seed for large reproductive efforts within a plant | 不需要植物个体之间的同步生产, 因此不能单独解释大年结实; 可能夸大了大小年间的变化 Does not require synchrony between individuals, hence cannot alone explain mast seeding; may exaggerate variation between high and low years ( | |
捕食者净化 Predator cleansing | 竹子的同步性死亡降低了以成年树叶为食的植食性动物的密度 The synchronized death of bamboos reduces densities of herbivores feeding on adult leaves ( | 群体开花可能是为了逃避捕食 Gregarious flowering may escape predation | 较少验证 Less empirical test | |
假说 Hypothesis | 描述 Description | 典型特征 Typical characteristics | 评论 Comment | |
大年抑制 Mast depression | 植物种子产量的周期性变化导致植食性动物种群的周期性波动 The cyclic population fluctuation of herbivores is caused by the cyclic seed crops of plants ( | 高种子产量的生产是以牺牲对植食性动物的化学防御为代价的 The production of high-seed crops is at the cost of chemical defense against herbivores | 主要用种子内化学防御物质的多少来解释植物与捕食者间的关系 The relationship between plants and predators is largely explained by the amount of chemical defenses in seeds | |
异型杂交 Outcrossing | 大年结实有利于杂交繁殖 Mast seeding facilitates outbreeding ( | 大量开花可减少近交衰退 Mast seeding may reduce inbreeding depression | 没有得到很好验证 Not well supported | |
竹子火烧循环 Bamboo fire cycle | 竹子等无性系种群大年盛花后同步死亡, 这助长了火灾的发生, 从而阻止了树木与竹子之间的竞争 Synchronized death of clonal populations such as bamboos after high flowering encourages fire, which prevents trees from out-competing the bamboo ( | 竹类等植物依赖林窗更新和扩散种群的一种适应对策 An adaptive strategy for plants such as bamboo that depends on forest gap regeneration and dispersal | 仅是一种推论, 需要实验验证 It is just a inference that needs to be tested experimentally | |
物候同步 Phenological synchrony | 大年结实是由个体的资源限制和环境变化引起的种群范围的花粉可利用性共同导致 The mast seeding is caused by both individual resources limitation and population-wide pollen availability due to environmental variation ( | 物候的作用导致植物同步开花从而增加授粉的有效性 Phenology leads to synchronous flowering of plants, increasing pollination effectiveness | 仅在山谷橡树中得到验证, 需更多实验验证 Only documented in the valley oak ( |
表1 解释大年结实现象的各种假说
Table 1 Hypotheses which have been proposed to explain the phenomenon of mast seeding
假说 Hypothesis | 描述 Description | 典型特征 Typical characteristics | 评论 Comment | |
---|---|---|---|---|
资源匹配(气候跟踪) Resource matching (Weather tracking) | 种子产量的变化是由于植物可利用资源的变化造成的 Seed crops vary in response to plant available resources variation ( | 在不改变营养生长或储备资源的情况下实现种子产量的变化 Variable seed crops achieved without switching resources from vegetative growth or reserves | 较早被提出, 早期采用反证法; 多数支持而少数不完全支持 Put forward earlier, and early adopt proof by contradiction ( | |
捕食者饱和 Predator satiation | 大量结实使捕食者饱和, 导致个体被捕食的比例降低 Large seed crops satiate seed predators, which thus lead to individuals experience lower proportion seed predation ( | 种子大年捕食率较低 Lower predation rate in high-seed years | 被多数研究支持; 不过, 取决于功能群的类型 Well supported by many studies ( | |
风媒授粉 Wind pollination | 风媒授粉植物同步大量开花可以获得更高的传粉效率 Wind-pollinated plants can achieve greater pollination efficiency through synchronized above-average flowering effort ( | 在大量开花年受精比例更高 Higher % fertilization in high flowering years | 很好地被验证; 然而, 部分实验结果是模棱两可的 Well documented ( | |
动物扩散(捕食者扩散) Animal dispersal (Predator dispersal) | 大年结实吸引更多扩散者, 因而刺激分散贮藏动物更多的扩散和贮藏种子 Large fruit production attract more seed dispersers, and result in wider dispersal and hoard by scater-hoarders ( | 种子大年扩散率更高或扩散更远 Higher seed dispersal rate or greater dispersal distances in high-seed years | 是否有利于持续繁殖取决于扩散者的功能反应; 部分结论支持, 部分反对 Whether favor constant reproduction, depending on functional response of dispersers; some evidences in support ( | |
环境预测 Environmental prediction | 植物可以“预知”最适气候条件而繁殖和建成 Plants can “predict” favorable climate conditions for reproduction and establishment ( | 种子或幼苗在大年结实时生存率更高 Higher seed or seedling survival in mast seeding | 部分证据支持, 部分反对 Some evidences in support ( | |
大种子 Large seed | 生产大种子增加了种子大年间的恢复时间 Selection for larger seed size increases the recovery time between large seed crops ( | 在种子较大的物种中, 种子数随年际变化更大 More variable seed number among years in related species with large seeds | 部分结论支持, 部分反对; 其他研究认为种子大小和种子数量存在权衡; 中等大小的种子具有更高的建苗率 Some support ( | |
动物传粉 Animal pollination | 大量开花更易于吸引动物传粉者, 从而提高传粉效率 Large flowering attract animal pollinators, thus enhance pollination efficiency ( | 在大量开花年受精比例更高 Higher % fertilization in high-flowering years | 一些分布稀少的植物个体同时产生大量的花, 更有助于种群数量的增加 A few sparsely distributed individuals that simultaneously produce a large number of flowers are more conducive to population growth ( | |
辅助成本 Accessory cost | 较高的固定繁殖成本有利于更少、更大规模的繁殖活动 Higher fixed reproductive costs favor fewer, larger reproductive episodes ( | 在植物内进行大量繁殖时, 每粒种子的辅助成本较低 Lower accessory costs per seed for large reproductive efforts within a plant | 不需要植物个体之间的同步生产, 因此不能单独解释大年结实; 可能夸大了大小年间的变化 Does not require synchrony between individuals, hence cannot alone explain mast seeding; may exaggerate variation between high and low years ( | |
捕食者净化 Predator cleansing | 竹子的同步性死亡降低了以成年树叶为食的植食性动物的密度 The synchronized death of bamboos reduces densities of herbivores feeding on adult leaves ( | 群体开花可能是为了逃避捕食 Gregarious flowering may escape predation | 较少验证 Less empirical test | |
假说 Hypothesis | 描述 Description | 典型特征 Typical characteristics | 评论 Comment | |
大年抑制 Mast depression | 植物种子产量的周期性变化导致植食性动物种群的周期性波动 The cyclic population fluctuation of herbivores is caused by the cyclic seed crops of plants ( | 高种子产量的生产是以牺牲对植食性动物的化学防御为代价的 The production of high-seed crops is at the cost of chemical defense against herbivores | 主要用种子内化学防御物质的多少来解释植物与捕食者间的关系 The relationship between plants and predators is largely explained by the amount of chemical defenses in seeds | |
异型杂交 Outcrossing | 大年结实有利于杂交繁殖 Mast seeding facilitates outbreeding ( | 大量开花可减少近交衰退 Mast seeding may reduce inbreeding depression | 没有得到很好验证 Not well supported | |
竹子火烧循环 Bamboo fire cycle | 竹子等无性系种群大年盛花后同步死亡, 这助长了火灾的发生, 从而阻止了树木与竹子之间的竞争 Synchronized death of clonal populations such as bamboos after high flowering encourages fire, which prevents trees from out-competing the bamboo ( | 竹类等植物依赖林窗更新和扩散种群的一种适应对策 An adaptive strategy for plants such as bamboo that depends on forest gap regeneration and dispersal | 仅是一种推论, 需要实验验证 It is just a inference that needs to be tested experimentally | |
物候同步 Phenological synchrony | 大年结实是由个体的资源限制和环境变化引起的种群范围的花粉可利用性共同导致 The mast seeding is caused by both individual resources limitation and population-wide pollen availability due to environmental variation ( | 物候的作用导致植物同步开花从而增加授粉的有效性 Phenology leads to synchronous flowering of plants, increasing pollination effectiveness | 仅在山谷橡树中得到验证, 需更多实验验证 Only documented in the valley oak ( |
图1 一个整合气候、资源和动植物互作的进化与生态过程的大年结实概念框架图
Fig. 1 A frame-diagram explaining the evolutionary and ecological processes of mast seeding by integrating the climate and resource and plant-animal interactions
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