植物大年结实及其与动物贮食行为之间的关系

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。     

种子特征和结实量对啮齿动物取食和扩散种子行为的影响

为了深入了解啮齿动物在不同种子丰富度条件下对不同大小和单宁含量种子的觅食行为策略及其与植物种群更新的关系,在宁夏六盘山区的华北落叶松人工林,研究了不同大小和单宁含量[0%Tannin(T)、2%T、8%T和15%T]的人工种子在模拟结实小年和结实大年对啮齿动物取食和扩散行为的影响.结果表明: 啮齿动物消耗种子速度在结实小年更快,结实大年的种子消耗速度相对缓慢. 种子就地取食率(ISPR)在不同结实年份间无显著差异,扩散后取食率(PRAD)在结实小年显著高于结实大年,但前者的扩散后贮藏率(HRAD)显著低于后者;种子扩散后的取食距离(PDAD)和贮藏距离(HDAD)在结实小年均显著大于结实大年.在结实小年,大种子的PDAD和HDAD均大于小种子,前者在不同大小种子间均差异显著,而后者仅在2%T和15%T的不同大小种子间差异显著;在结实大年,除0%T外的其他单宁含量种子的PDAD和HDAD在不同大小种子间均差异显著.ISPR在中等单宁含量种子最大,高单宁含量种子最小;PRAD分别在结实小年的高单宁含量种子和结实大年的无单宁种子最大;不论在结实大年还是结实小年,HRAD均在高单宁含量种子最大,中等单宁含量种子最小.这说明结实大年可延缓啮齿动物对种子的消耗速率,提高种子的HRAD,但种子扩散距离减小;啮齿动物在结实大年和小年均表现出对大种子的扩散偏好,且大种子被扩散的距离更远;啮齿动物在不同结实年份均偏好于就地取食中等单宁含量种子,而扩散高单宁含量种子.     

红松结实量与松果可利用量对松鼠和花鼠贮食行为的影响

植物通过每隔几年产生大量种子的丰年策略提高贮食动物传播种子效率,但人为采摘活动降低了种子的可利用量,从而影响贮食动物行为及种群动态。为研究红松球果采摘如何通过降低松果可利用量而影响贮食动物行为,我们基于黑龙江凉水自然保护区2003-2012年红松结实量和采摘量的变化,比较分析了结实大小年间松鼠贮点大小、贮点深度、贮食密度和贮藏量、松鼠花鼠种群以及2010年和2011年花鼠洞穴贮藏量的差异。结果显示：结实大年松鼠的平均贮点大小显著高于小年,大贮点比例增加,贮食密度和贮藏量也明显高于结实小年,但随着结实量的增大,球果采摘量增加,使松果可利用量减少,由此结实大年2011年松鼠贮食密度并未随结实量增加而增加,反而低于2003年和2008年,松鼠遇见率也没有在该结实大年有所增长。而花鼠种群和洞穴松籽贮藏量在结实大年和小年间没有显著差异。研究表明红松球果采摘对松果可利用量和松鼠贮食行为有较大影响,应合理确定大年的采摘量以保证贮食动物的食物丰度,维系红松生态系统健康。     

林木大年结实特征及其影响因素研究进展

大年结实是树木普遍的繁殖行为,在种群水平表现出较高的年际变化,存在不规律的间隔期。这种结实规律有利于调节捕食者种群数量,提高授粉效率,促进种子散布和降低种子捕食者的影响,是一种进化的生殖策略,但其对生产上种实高产稳产非常不利。本文综述了大年结实的特点以及大年结实对树木营养生长、资源消耗的影响,分析了影响大年结实的主要因素(碳水化合物、矿质养分、激素水平、气候因子等),最后提出了我国大年结实研究应该努力的方向。     

被子植物地下结实和地上/下两型结实的生态适应意义

地下结实和地上/下两型结实是被子植物两类独特的结实方式, 多发生在陆生和草本植物中, 主要生长在缺少水分或光照、土壤扰动频繁及环境波动较为剧烈的生境中。两种结实方式不仅是植物适应性进化的重要方面, 也是选择性进化的产物。其中, 地下结实对于植物在母株附近适宜微环境中保存后代、在极端环境下保持种子活力、逃避地面动物取食和火灾伤害以及延长果实发育时间等方面, 地上/下两型结实对于减少同胞子代及种群内竞争、维持和扩大种群以及提高物种的适应力和进化可塑性等方面, 都具有重要的生态适应意义, 是植物抵御不利生物与非生物环境的两类重要防御策略。但两种结实方式同时也存在着限制果实与种子扩散、影响基因传递与种群遗传结构、加大种群隔离以及提高繁殖代价等进化限制, 对物种的分布、种群增长、迁移、适合度和生活史进化等具有重要影响。目前, 地下和地上/下两型结实现象分别在大约24科57属和13科34属中进行了报道, 其中在菊科、十字花科、豆科和玄参科等类群中两种结实现象同时存在。从系统发育看, 地下结实在木兰分支、单子叶植物分支及双子叶植物分支中均存在, 而地上/下两型结实仅出现在单子叶植物分支和双子叶植物分支中, 在被子植物基部类群(ANITA类群)中两种结实方式均不存在。该文对植物地下和地上/下两型结实的类型、系统进化、繁殖特性和扩散对策进行了介绍, 并对其生态适应意义进行了总结, 以期为深入研究植物结实的进化策略提供参考。     

蒙古莸的开花物候与生殖特征

通过对蒙古莸自然种群和人工种群的开花物候进程观察,统计不同种群的结实率,分析开花物候对其生殖成功的影响。结果表明:(1)蒙古莸花期在7月末至9月中旬,人工种群较自然种群的开花物候明显提前,其中始花期提早4～5d,但两个种群的花期持续天数基本一致。(2)蒙古莸个体间具有较高的开花同步性,不同种群蒙古莸植株的开花同步性指数均大于0.80,且自然种群略高;晴好天气下绝大多数花朵都集中在上午10:00以前开放,阴雨天则延迟开放或者不开放,这种"集中开花模式"增加了传粉者的访问频率,是蒙古莸提高生态适应性、保障其生殖成功的一种表现。(3)开花物候指数与结实数之间的相关分析表明,花序水平上,开花数及花期持续时间与结实数呈极显著正相关关系。     

草地植物种群繁殖对策研究

植物的繁殖包括有性繁殖和无性繁殖两大类型,克隆繁殖是一种较为特殊的营养繁殖方式。本文综述了草地种子植物的生殖分配及生殖投资,克隆生长以及放牧对草地植物种群繁殖的影响。植物种群生物量、能量和养分生殖分配是植物种群生殖分配的重要内容,不同植物在结实期营养元素及能量的配置上有着显著的区别,这可能是植物在长期进化过程中形成的生殖对策,是适应环境的结果。在种群水平上,中等强度以上的放牧干扰有利于植物的克隆生长,但有性生殖减弱。草原植物发达的营养繁殖或克隆生长方式是对放牧的适应性对策。     

东北草原羊草种群结实特性与气候年变化的关系

对３个固定样地的羊草（Ｌｅｙｍｕｓ ｃｈｉｎｅｎｓｉｓ（Ｔｒｉｎ．）Ｔｚｖｅｌ．）种群连续１２￣１６年的调查和测定及其与不同生长发育阶段气候因子的相关分析,表明羊草种群结实数量和籽实重量均与形成这些性状的生育期内,及其以前各生育期不同阶段的光照时间、积温、降水量有着一定程度的相关关系。从分蘖株的营养生长到生殖生长,包括开花、授粉、受精、胚珠发育为种子形成的整个过程中,较低的温度和生长季前期较多的光     

不同生态环境条件沙生植物沙鞭的结实研究

为了阐明不同生态环境条件下沙生植物沙鞭的结实规律,该文对沙鞭137个种群结实情况进行了实地观察,发现沙鞭种子的结实情况可被划分为无种子、种子饱满和种子不饱满3种类型;在此基础上,该文采用聚类分析、Kruskal-Wallis检验和典范对应分析(CCA)等方法探究沙鞭种群结实情况与22个地理气候因子的相关性。结果表明:(1)沙鞭137个种群按照地理气候因子不同聚为3个组;(2) Kruskal-Wallis检验显示沙鞭3个组间种子结实情况差异不显著(P=0.269),即沙鞭种群间种子结实与其所处的地理气候因子无直接相关性;(3)典范对应分析(CCA)表明沙鞭种群间种子结实情况差异也不显著(P0.05),但地理气候因子与种子饱满度以及无种子特征具有显著相关性,其中海拔和降雨因子(bio12-bio19)与种子饱满度呈正相关,而经纬度和温度因子(bio4,bio7)与种子饱满度呈负相关,无种子特征仅与最湿季平均温度(bio8)呈正相关。地理气候因子对沙鞭天然种群有性繁殖(有种子)重要性高于无性繁殖(无种子),表明制约沙生植物沙鞭有性繁殖的环境因子复杂,其无性繁殖可能是种群数量稳定的适应性表现。     

不同生境与花部特征对巫山淫羊藿结实特性的影响

对不同生境中巫山淫羊藿结实特性进行比较研究。结果表明:(1)三个种群在繁殖时期中不同花部特征的植株繁殖投资差异不大,如每株分枝数、每株花序数和每株开花数差异均不显著;(2)三个种群中每株结实数、每株结实率、每花序结实数在不同花部特征中差异显著,PLS和SAA要显著大于PESS;(3)三种花部特征在不同生境中单株分枝数和单株花序数均无显著差异。在不同花部特征中种群1开花数量相较少,因而在结实方面均值均要小于种群2和种群3;(4)不同花部特征的果实饱满种子数、败育数和败育率均具有显著差异,而种子数差异不大,但在种群间均无显著的差异。说明种子的多少更多地受到花部特征的影响。总之,巫山淫羊藿的结实特性受到生境和花部特征的双重影响,其中花型比生境对巫山淫羊藿结实特性的影响更为明显。     

Measuring mast seeding behavior: relationships among population variation,individual variation and synchrony

Mast seeding, or masting, is the variable production of flowers, seeds, or fruit across years more or less synchronously by individuals within a population. A critical issue is the extent to which temporal variation in seed production over a collection of individuals can be viewed as arising from a combination of individual variation and synchrony among individuals. Studies of masting typically quantify such variation in terms of the coefficient of variation (CV). In this paper we examine mathematically how the population CV relates to the mean individual CV and synchrony, concluding that the relationship is a complex one which cannot isolate an overall measure of synchrony, and involves additional factors, principally the number of plants sampled and the mean productivity per plant. Our development suggests some simple approximate relationships of population CV to individual variability, synchrony and the number of individuals. These were found to fit quite well when applied to data from 59 studies which included seed production at the individual level.     

Spatial dynamics of specialist seed predators on synchronized and intermittent seed production of host plants

Masting, the synchronized and intermittent seed production by plant populations, provides highly variable food resources for specialist seed predators. Such a reproductive mode helps minimize seed losses through predator satiation and extinction of seed predator populations. The seed predators can buffer the resource variation through dispersal or extended diapause. We developed a spatially explicit resource-consumer model to understand the effect of masting on specialist seed predators. The masting dynamics were assumed to follow a resource-based model for plant reproduction, and the population dynamics of the predator were represented by a spatially extended Nicholson-Bailey model. The resultant model demonstrated that when host plants reproduce intermittently, seed predator populations go locally extinct, but global persistence of the predator is facilitated by dispersal or extended diapause. Global extinction of the predator resulted when the intermittent reproduction is highly synchronized among plants. An approximate invasion criterion for the predators showed that negative lag-1 autocorrelation in seeding reduces invasibility, and positive lag-1 cross-correlation enhances invasibility. Spatial synchronization in seeding at local scale caused by pollen coupling (or climate forcing) further prevented invasion of the predators. If the predators employed extended diapause, extremely high temporal variability in reproduction was required for plants to evade the predators.     

Pollen limitation as a main driver of fruiting dynamics in oak populations

In many perennial wind‐pollinated plants, the dynamics of seed production is commonly known to be highly fluctuating from year to year and synchronised among individuals within populations. The proximate causes of such seeding dynamics, called masting, are still poorly understood in oak species that are widespread in the northern hemisphere, and whose fruiting dynamics dramatically impacts forest regeneration and biodiversity. Combining long‐term surveys of oak airborne pollen amount and acorn production over large‐scale field networks in temperate areas, and a mechanistic modelling approach, we found that the pollen dynamics is the key driver of oak masting. Mechanisms at play involved both internal resource allocation to pollen production synchronised among trees and spring weather conditions affecting the amount of airborne pollen available for reproduction. The sensitivity of airborne pollen to weather conditions might make oak masting and its ecological consequences highly sensitive to climate change.     

Synchronicity,periodicity and bimodality in inter‐annual tree seed production along an elevation gradient

Distinctive inter‐annual patterns of tree seed production can include spatial synchronicity, periodicity, and high variability among individuals within a population. Synchronicity and high variability are now commonly used to define mast seeding, with ‘strict’ mast seeding further distinguished by annual seed production that is either often large or nil and thus bimodal. Here we test for synchronicity, periodicity, and bimodality using 43 years of annual total and viable seed counts, along four transect lines, sampling an elevation gradient (480–1340 m) in a monospecific New Zealand mountain beech Nothofagus solandri var. cliffortioides forest. We expect most support for periodicity and bimodality at relatively high elevation sites that are most resource limited and known to have the greatest variability in seed production. While there was weak evidence for differences in viable seed counts along the elevation gradient, this was not the case for total seed counts. A significant year‐effect on seed counts provided some evidence for synchronization, but the correlations of total or viable seed counts rapidly declined (from 0.96 to 0.58) with increasing elevation difference among transect lines. Although we detected a seven‐year periodicity in total and viable seed counts at each elevation, we also detected other period lengths at most elevations. We did not find evidence for our expectation of increased period length and bimodality in relatively unproductive high elevation mountain beech forests because they would take more time to recover from seeding events. As a consequence, if resource limitation is an important driver of mountain beech seed production its influence must be strongly regulated by other factors to determine the distinctive characteristics (periodicity and synchronicity but not bimodality) of inter‐annual variation in seed production.     

Flowering phenology of <Emphasis Type="Italic">Ulex europaeus</Emphasis>: ecological consequences of variation within and among populations

Reproductive phenology of gorse (Ulex europaeus L., Genisteae, Fabaceae) is unusual in that the onset and duration of flowering vary greatly among individuals within populations: some plants initiate flowering in autumn or winter and continue flowering through spring, others initiate flowering in early spring. To understand the origin of this diversity and its ecological consequences, we investigated flowering phenology of randomly sampled individuals from five different natural populations in Brittany (France). Reproductive success was evaluated for individuals with contrasting flowering patterns, from 16 natural populations. Flower production, pod production, seed production and seed predation were estimated. Plants initiating flowering in spring produced larger numbers of flowers and pods over a shorter period than plants flowering from winter to spring, which produced few flowers and pods at a time but over a longer period. Pod production of long-flowering plants did not differ significantly between winter and spring, but their pods were more intensively attacked by seed predators in spring than in winter. We discuss our results in relation to biotic and abiotic parameters. We postulate that long-flowering can be interpreted as a bet-hedging strategy, spreading the risk of pod failure (rotting or freezing) in winter and of seed predation in spring.     

EFFECTS OF POSTDISPERSAL SEED PREDATION ON SPATIAL INEQUALITY AND SIZE VARIABILITY IN AN ANNUAL PLANT,ERODIUM CICUTARIUM (GERANIACEAE)

By decreasing seed density, ants introduced into flats of uniformly sown seeds of Erodium cicutarium (Geraniaceae) created differences in the neighbor-free area available to individual plants. The changes in spatial patterns brought about by the ants were greater when a higher proportion of seeds was removed but were independent of initial seed density. These spatial changes and differences in seed density were examined for their effects on plant size and reproduction. Gini values were calculated to determine inequalities. As the inequality in space among individual plants increased, the variation in final biomass increased. The number of individuals reproducing was constant among treatments, and yet seed production per plant was significantly greater for populations in which the spatial pattern was influenced by seed predation. The decrease in density and changed spatial pattern, due to previous seed predation, resulted in a few individuals having much more space than others and consequently producing many more seeds. The increase in reproductive effort per flat was much greater than could be explained by the changing density alone. Our experiment demonstrates that spatial inequality, such as that generated by seed predators, can be more important than density in generating size inequalities in plant populations. This result can profoundly alter the competitive interactions between plants and determine which plants produce seed for the next generation.     

Mast seeding in Abies balsamea, Acer saccharum and Betula alleghaniensis in an old growth, cold temperate forest of north-eastern North America

1 Several hypotheses have been put forward to explain the phenomenon of masting or mast seeding, i.e. the supra-annual, periodic production of a large number of seeds in long-lived plants. Some of these hypotheses deal with the proximate causes of masting (e.g. the climate hypothesis) but others are concerned mostly with ultimate, evolutionary explanations (e.g. the pollination efficiency hypothesis).
2 The seed production of three tree species, Abies balsamea , Acer saccharum and Betula alleghaniensis , was followed over a 7-year period in an old-growth, cold temperate forest of north-eastern North America. The main objectives were to determine the extent of interannual variations in seed production, to investigate the relationship between viable and potential seed crop and crop efficiency, and to explore the effects of climate on seed production.
3 Potential and viable seed production varied significantly among years for all three species. However, the timing of dispersal remained the same regardless of the level of seed production.
4 Seed rain was spatially less heterogeneous in years of high seed production, suggesting that most trees were reproducing in such years.
5 Over the 7-year period, there was a significant concordance among species in their viable seed crop and crop efficiency, but not in their potential seed crop. Crop efficiency was positively correlated to potential seed crop for Abies and Betula , but not for Acer .
6 High seed production was related to warm, dry conditions in the spring of the previous year (i.e. at reproductive bud initiation) but to a moist summer in the year of seed maturation.
7 Masting in these three species thus appears to be controlled by several factors, including climate and pollination efficiency.     

Effects of mast seeding and rodent abundance on seed predation and dispersal of <Emphasis Type="Italic">Quercus aliena</Emphasis> (Fagaceae) in Qinling Mountains,Central China

Rodents act as seed predators and dispersers and play an important role in the regeneration of plants. Seed production and rodent abundance may influence caching rodents’ decision to consume or cache seeds. We studied how did seed production and rodent abundance co-influence seed predation and dispersal by rodents in the mast seed years (2011 and 2013) and non-mast seed years (2012 and 2014) in the Qinling Mountains of Shaanxi Province, Central China. We found that: (1) The seed removal rates were much faster in the non-mast seed years than in the mast seed years. (2) Although the total number of seeds consumed (including eaten in situ and after removal) was higher in the non-mast seed years than in the mast seed years, no significant differences were found in the seeds that were eaten in situ and after removal among the 4 years. (3) Significant differences were observed in seeds that were cached among the 4 years, and more seeds were cached in the non-mast seed years than in the mast seed years. (4) The primary and secondary seed dispersal distances (including cached and eaten) were all longer in the non-mast seed years than in the mast seed years. Overall, these results indicate that non-mast seeding with low per capita seed availability could increase both seeds consumption and caching by rodents. Our results are partially supporting the predator satiation hypothesis.     

Tree-to-tree variation in seed size and its consequences for seed dispersal versus predation by rodents

Individual variation in seed size and seed production is high in many plant species. How does this variation affect seed-dispersing animals and, in turn, the fitness of individual plants? In this study, we first surveyed intraspecific variation in seed mass and production in a population of a Chinese white pine, Pinus armandii. For 134 target trees investigated in 2012, there was very high variation in seed size, with mean seed mass varying among trees almost tenfold, from 0.038 to 0.361 g. Furthermore, 30 of the 134 trees produced seeds 2 years later, and for these individuals there was a correlation in seed mass of 0.59 between years, implying consistent differences among individuals. For a subset of 67 trees, we monitored the foraging preferences of scatter-hoarding rodents on a total of 15,301 seeds: 8380 were ignored, 3184 were eaten in situ, 2651 were eaten after being cached, and 395 were successfully dispersed (cached and left intact). At the scale of individual seeds, seed mass affected almost every decision that rodents made to eat, remove, and cache individual seeds. At the level of individual trees, larger seeds had increased probabilities of both predation and successful dispersal: the effects of mean seed size on costs (predation) and benefits (caching) balanced out. Thus, despite seed size affecting rodent decisions, variation among trees in dispersal success associated with mean seed size was small once seeds were harvested. This might explain, at least in part, the maintenance of high variation in mean seed mass among tree individuals.     

Pollen viability and limitation of seed production in a population of the circumpolar cushion plant, Silene acaulis (Caryophyllaceae)

Pollen viability among genders and limitation of female seed production in a natural trioecious population of the circumpolar cushion plant Silene acaulis was examined. Pollen viability was estimated by an in vitro pollen germination experiment. Both male and hermaphrodite flowers displayed large variation in pollen viability (0–53% in hermaphrodite and 0–54% in male flowers). There was a significant difference between genders in pollen viability: male plants had on average higher pollen viability than hermaphrodite plants. Resource and pollen limitation of seed production was studied by an experiment consisting of three treatments; (I) hand-pollination and removal of all other flowers on the cushion, (II) hand-pollination without removal of other flowers, and (III) open pollination without removal of flowers. Hand-pollination increased seed production, whereas removal of flowers had no effect on seed production. Abortion of pollinated ovules during seed development and seed mass did not differ among treatments. To control for effect of fruit number on seed production, data from naturally pollinated individuals was used. There was a positive correlation between both total number of seeds and fruit number, mean seed number per fruit and fruit number, respectively. These results indicate that seed production of 5. acaulis is mainly limited by pollen availability whereas resource competition between fruits is not important as a limiting factor. The possible role of male quality differences between genders and pollen limitation of seed production for maintenance of trioecious reproductive systems is discussed.