植食性哺乳动物能量收益函数模型的预测性及适用性

以东方田鼠喜食的白三叶叶片作为食物,在保持叶片生物量不变的条件下,改变叶片大小,配置东方田鼠觅食的各类食物大小异质斑块,测定东方田鼠觅食的行为。通过比较几种植食性哺乳动物能量收益函数模型的预测性,评价其适用性。结果发现,没有检测到东方田鼠食物摄入量动态呈S型能量收益函数增长。线性函数模型能准确地预测东方田鼠在中、小型食物斑块的停留时间;在大型食物斑块,尽管,分段线性函数及渐进函数均能很好地拟合东方田鼠的食物摄入量动态,但仅分段线性函数模型能准确地预测其停留时间。线性函数模型及分段线性函数模型是在功能反应机制模型-口量模型基础上建立的,反映了调节摄入率动态的机制,为机制性模型。因此,此2种模型是在口量及摄食站尺度上,探讨动物在食物斑块的能量收益动态及停留时间;而渐进函数模型及S型函数模型均为实验性模型,是在斑块尺度上预测动物能量收益动态及停留时间,未能反映动物摄入率动态,故其预测效果较差。由于此4种模型均未考虑动物在食物斑块搜寻食物及非觅食活动如警觉和逃跑等花费的时间,因而,限制了模型的广泛应用。建议,发展新的模型,促进觅食生态学斑块模型理论研究的深入发展。     

行为生态学(二):最优觅食行为(1)

动物觅食时经常面临着各种不同的选择,如猎取什么食物?到哪里去猎食?采取怎样的猎食路线?自然选择总是使动物的觅食效率尽可能地加以改进,因为只要捕食者能够借助于更有效的觅食,使其存活和繁殖成功的机会增加,自然选择就会对它有利。这个道理对山雀属鸟类来说是很明显的,这些食虫小鸟在冬季白天必须每隔三秒钟就捕食一只昆虫才能维持自己的生存(J.A.Gibb,1960)。     

视野受阻对东方田鼠觅食行为的影响

植食性小型哺乳动物的觅食行为是对特定生境的适应性产物。食物斑块周边植被对动物视野遮挡是否通过作用于其觅食活动中的警觉而影响摄入率。采用新鲜白三叶叶片构建东方田鼠密集均质食物斑块,以牛皮纸模拟食物斑块周边植被遮挡田鼠视野,测定其在食物斑块上的觅食行为序列过程及行为参数,检验食物斑块周边植被高度对东方田鼠摄入率的影响。结果发现,个体在不同程度视野受阻条件下食物摄入率无显著差异。分析觅食行为参数动态发现,在不同视野受阻条件下,个体能通过调整各采食回合内警觉行为动作的发生频次和持续时间,维持觅食回合内总的觅食中断时间的稳定,进而保证进食时间的稳定。东方田鼠在不同程度的视野遮挡条件下均能通过行为变异和优化使摄入率保持稳定。结果亦充分说明,东方田鼠在警觉强度上的变化不能反映觅食中断所带来的食物收益减损的代价,但觅食活动中各警觉动作的持续时间的变异却能够明确指示个体摄入率的动态变化,因此,以觅食活动中警觉引起的觅食中断时间代价为线索,检验其对摄入率的影响,是评价植食性小型哺乳动物在不同生境觅食适应性策略的有效的方法。     

行为生态学(五):动物的觅食技巧

动物的觅食行为包括对食物的搜寻、捕捉和加工处理三个方面。动物觅食的搜寻方式、捕食效率和处理技能都对动物的食性范围有直接影响。但是这三种觅食技能的相对重要性却依食物的特性而有所不同,以叶栖昆虫为食的小型鸟类就是这方面的一个典型事例,这些鸟类专门以小型和隐蔽的猎物为食,所以它们特别善于搜寻,而不太善于追捕和对付那些比较活跃的动物。下面就动物觅食行为的三种技能分别作些简要介绍。     

捕食风险对东方田鼠功能反应格局的作用

植食性哺乳动物对食物斑块的选择和利用不仅取决于食物的可利用性,且与觅食环境潜存的各种风险紧密关联。捕食风险是否通过作用于动物觅食活动中的警觉影响其功能反应格局。在新鲜白三叶叶片构成的各类食物密集斑块上,测定东方田鼠觅食行为,建立功能反应模型,检验捕食风险对其功能反应格局的作用。结果发现,捕食风险能显著地延长东方田鼠的觅食决定时间,但其摄入率保持稳定,功能反应构型亦未发生改变,仍为Ⅱ型功能反应;除了对照组个体的采食时间随叶片大小增大无明显变动规律外,处理组个体的采食时间及对照组和处理组个体的处理时间、觅食中断时间均随叶片大小及口量的增大呈线性增高趋势,处理组个体的觅食中断时间明显大于对照组个体的;对照组和处理组个体的采食率均随叶片大小及口量呈非线性渐近递减趋势,但处理组个体的采食率较对照组个体的略有降低。结果揭示,在捕食风险压力下,虽然上述觅食参数变异能潜在地降低摄入率,但个体能通过改变觅食活动中各种警觉行为动作如降低嗅闻和静听监视动作的发生频次,增大视觉监视动作比重,以此缓冲捕食风险压力,维持摄入率。摄入率测定值与模型预测值的线性回归极显著,表明,功能反应模型具有良好的预测性。在可利用植物密集斑块,动物觅食活动中的警觉能缓冲捕食风险压力;动物摄入率是由植物大小调控的口量决定的,且受采食与处理食物竞争及觅食中断的制约;其功能反应仍属Ⅱ型功能反应。     

植食性哺乳动物觅食的功能反应及其模型

章主要介绍植食性哺乳动物觅食功能反应与模型的研究进展。植食性哺乳动物觅食的摄入率与其食物可利用性的功能反应是动物觅食生态学过程的基础。可利用植物的生物量密度、植物密度、植物大小、以及动物觅食的口量是影响动物觅食功能反应的潜在变量集。这些变量的差异导致动物功能反应格局的复杂化。生物量密度和植物密度对动物摄入率无明显影响,而植物大小对动物摄入率则有显影响。有食物密集的斑块条件下,以植物大小代替动物     

植食性哺乳动物食物选择的生态学意义及行为机制

综述了植食性哺乳动物食物选择行为机制的研究进展。植食性动物的食物选择是觅食生态学研究最为活跃的领域,动物食物选择对策主要有营养选择、植物次生化合物假设、营养平衡假设、最优觅食理论和条件性气味回避假设。动物学习食物选择的行为过程中,觅食个体能通过其认知过程和感知过程来处理食物信息、学习选择食物项目是一种可塑性行为方式,有利于动物获得适应栖息地的行为模式。幼体通过社会学习自母体学习的觅食经验,在幼体一生的食物选择中均具有重要的作用。在环境条件相对稳定条件下,幼体可模仿成体的食物选择模式;在环境剧烈变化的条件下,动物通过试错学习选择的食物项目可能较模仿成体所摄取的食物项目营养含量更丰富。     

植食性哺乳动物能量收益增长减速机制的检验

能量收益函数描述了植食性哺乳动物在植物斑块的食物摄入量与在斑块停留时间的函数关系,为觅食生态学理论的重要组成部分。在新鲜白三叶叶片构成的各类叶片斑块上,测定东方田鼠的觅食行为,建立其能量收益函数模型,分析植食性哺乳动物能量收益增长减速的机制。研究结果表明,东方田鼠觅食大叶片时,叶片干物质收益与停留时间呈非线性渐进函数关系,能量收益函数为渐进函数;觅食中、小型叶片时,叶片干物质收益与停留时间呈线性函数关系,能量收益函数为线性函数;没有检测到东方田鼠的能量收益动态呈分段线性函数或S型函数增长。东方田鼠在大、中型叶片斑块觅食时,随停留时间的增大,口量呈线性或指数递减,而处理时间则呈线性或指数递增,采食时间、间隔时间及咀嚼频次保持相对稳定,瞬时摄入率呈减小趋势;东方田鼠在小叶片斑块觅食时,觅食行为参数口量、摄入率、采食时间、间隔时间、处理时间及咀嚼频次均保持相对稳定。研究结果充分验证了,植食性哺乳动物在消费植被过程中,大型可利用性植物减少,受植物大小调控的动物口量减小,处理时间增加,引起瞬时摄入率降低,导致其能量收益增长减速的假说。     

社群学习对植食性鸟类和哺乳动物觅食行为的作用

社群学习是动物的一种可塑性行为表现型式。综述了社群学习对植食性鸟类和哺乳动物觅食行为的作用,并述评了其学习机制。社群同伴对动物个体觅食地点、时间和取食方式均有影响,母体摄食的食物信息可通过胎盘和乳汁显著影响幼体的食物选择。动物通过观察学习、嗅闻学习以及味觉厌恶学习,不仅能更快找到食物资源,提高觅食效率,而且能有效降低中毒与被捕食的风险,从而提高其适合度。     

动物觅食行为对捕食风险的反应

动物进行任何活动时均面临被捕食的风险 ,分析捕食风险与猎物觅食行为的关系 ,有助于揭示捕食者与猎物的协同进化机制。捕食风险具有限制或调节猎物种群数量的功能。在进化时间内 ,对猎物形态和行为特征的进化是潜在的选择压力之一 ,可利用环境因子作为信息源估测食物可利用性和捕食风险大小的动物 ,具有更大的适合度。信息源可分为包括视觉的、听觉的和化学的。动物进行觅食活动时 ,依据信息源的变化确定环境中捕食风险的大小 ,并根据自身的质量在捕食风险的大小之间做出权衡 ,通过食物选择、活动格局和栖息地利用等行为的变化降低捕食风险     

Evolutionary models of metabolism, behaviour and personality

I explore the relationship between metabolism and personality by establishing how selection acts on metabolic rate and risk-taking in the context of a trade-off between energy and predation. Using a simple time budget model, I show that a high resting metabolic rate is not necessarily associated with a high daily energy expenditure. The metabolic rate that minimizes the time spent foraging does not maximize the net gain rate while foraging, and it is not always advantageous for animals to have a higher metabolic rate when food availability is high. A model based on minimizing the ratio of mortality rate to net gain rate is used to determine how a willingness to take risks should be correlated with metabolic rate. My results establish that it is not always advantageous for animals to take greater risks when metabolic rate is high. When foraging intensity and metabolic rate coevolve, I show that in a particular case different combinations of foraging intensity and metabolic rate can have equal fitness.     

Optimal foraging by deposit-feeding invertebrates: Roles of particle size and organic coating

Summary Feeding experiments were conducted on marine, deposit-feeding benthic invertebrates to test the predictions of an optimal foraging model. Food item selection based on sediment particle size and presence or absence of an organic coating on particles was investigated. Animals displaying a wide range of feeding mechanisms were studied in particle size-selection experiments using artificial sediment of closely controlled size composition. Nine of 10 species from 4 phyla ingested smaller particles in greater proportions than the particles were present in the sediment. In experiments where animals fed on a mixture of two particle types, one with and one without a surface protein coating, 6 of 7 species from 3 phyla ingested preferentially the protein-coated beads. While these trends of selection of smaller particles and protein-coated particles follow qualitatively the predictions of the optimal foraging model, the animals did not ingest exclusively the preferred particle types. Mechanics of particle handling rather than behavioral responses to particle characteristics appear to offer the better explanation for the observed selection patterns. In particular, the results support strongly the recently proposed role of mucous adhesion in particle selection by deposit feeders.These and other results from studies of deposit feeders suggest that factors in addition to food item selection must be considered when testing the assumptions and predictions of optimal foraging theory. Specifically, feeding energetics are also affected by postfood-selection processes such as variation of ingestion rate. Furthermore, the effects of abiotic environmental factors on foraging behavior cannot be overlooked in evaluating the optimality of foraging behavior; variable water velocity affected differently the particle size selectivity of 3 sympatric polychaete species in these studies.Contribution No. 1239 from the School of Oceanography, University of Washington     

The constraints of digestive rate: An alternative model of diet selection

Summary A general model is developed to predict diet selection when digestive capacity is limited and when food items differ in digestibility and digestive turnover time. Under these conditions, in order to maximize the rate of energy acquisition, animals should maximize digestive rate, by selecting food with high digestibility and rapid passage through the digestive tract. They should spend as much time as they have available to search for this food. These and other predictions differ from those of the widely used Contingency Model, which maximizes the rate of ingestion of energy. Many experiments in the literature have not discriminated between predictions of these two models. Moreover, clarification of the conditions under which these two general models apply leads to a new perspective on the diversity of foraging behavior in animals.     

Tracking a fluctuating environment: a study of sampling

The problem of how animals keep track of unpredictable changes in the profitability of foraging sites was studied. An optimality model was used to predict the frequency with which a forager should sample a foraging site in which the probability of reward fluctuates randomly between high and low. The alternative foraging site is stable and offers an intermediate probability of reward. The model was tested with pigeons in a shuttle-box the two ends of which represented the two foraging sites. The pigeons succeeded in tracking the changes in the fluctuating site and the payoff attained was close to the optimum. Variations in the frequency of sampling between experimental treatments were in qualitative agreement with the model for some treatments but not others. The quantitative details of sampling behaviour were not as predicted by the optimality model, but many features could be accounted for by a mechanistic model of choice. The pigeons' choice rule, although different from that of the optimality model, is one that produces near-optimal payoffs under the conditions of this experiment.     

An economic model of the limits to foraging range in central place foragers with numerical solutions for bumblebees

1. A model is described that evaluates the maximum economic foraging range in central place foragers by using optimality criteria to discriminate between foraging sites at different distances from the forager's central place. 2. The basic model can be varied to suit foragers that optimise either their rate of net energy uptake or their foraging efficiency. 3. The model requires specification of the time and energy budgets of travel and foraging, and of the rewards obtainable at potential foraging sites. 4. The specific case of bumblebees, whose foraging ranges are poorly known, is considered. 5. Numerical solutions of the model for parameter values that represent bumblebees and their forage predict economic foraging ranges exceeding several kilometres. The model demonstrates that economics alone can explain extensive flight ranges in bees.     

Structural complexity and fish body size interactively affect habitat optimality

Predator–prey interactions are strongly influenced by habitat structure, particularly in coastal marine habitats such as seagrasses in which structural complexity (SC) may vary over small spatial scales. For seagrass mesopredators such as juvenile fishes, optimality models predict that fitness will be maximized at levels of SC that enhance foraging but minimize predation risk, both of which are functions of body size. We tested the hypothesis that in eelgrass (Zostera marina) habitat, optimal SC for juvenile giant kelpfish (Heterostichus rostratus), an abundant eelgrass mesopredator in southern California, changes through ontogeny. To do this, we quantified eelgrass SC effects on habitat associations, relative predation risk, and foraging efficiency for three size classes of juvenile giant kelpfish. We found that habitat selection differed with fish size: small fish selected dense eelgrass, whereas larger fish selected sparse eelgrass. Small kelpfish experienced the lowest relative predation risk in dense eelgrass but also had higher foraging efficiency in dense eelgrass, suggesting that dense eelgrass is selected by these fish because it minimizes risk and maximizes potential for growth. Surprisingly, larger kelpfish did not experience lower predation risk than small kelpfish. However, larger kelpfish experienced higher foraging efficiency in sparse eelgrass vs. dense eelgrass, suggesting that they select sparse eelgrass to maximize foraging efficiency. Our study highlights that trade-offs between predation risk and foraging can occur within a single habitat type, that studies should consider how habitat value changes through ontogeny, and that seagrass habitat value may be maximal when within-patch variability in SC is high.     

Some effects of energy costs on foraging strategies

We consider the effect of including energy costs on the optimal strategy for animals exploiting a depleting food resource. In the context of central place foraging this leads to the problem of what load size should be brought back to the central place. Two strategies are discussed: (i) maximize gross rate of energy delivery and (ii) maximize net rate of energy delivery. The optimal load size (or optimal patch time) for net maximizers is not always larger than for gross maximizers, as has been claimed. Instead, the difference in optimal load size has the same sign as the difference between metabolic rates of travelling and foraging. We point out that the influence of costs has not always been correctly incorporated in experimental tests of the theory.     

Feeding and metabolic compensations in response to different foraging costs

How energetic cost of locomotion affects foraging decisions, and its metabolic consequences are poorly understood. In several groups of animals, including hermit crabs, exploratory walking enhances the efficiency of foraging by increasing the probability of finding more and better food items; however, the net gain of energy will only be enhanced if the costs of walking are lower than the benefits of enhanced food acquisition. In hermit crabs, the cost of walking increases with the mass of the shell type occupied. Thus, we expected that hermit crabs should adjust their foraging strategy to the cost of movement in different shells. We assessed the foraging, the quantity and quality of food intake, and the energetic cost of maintenance of hermit crabs paying different costs of foraging in the wild. The exploratory walking negatively correlated with shell mass, showing that hermit crabs use different foraging strategies in response to the expenditure required to move. Hermit crabs deal with high energetic costs of foraging in heavy shells by reduces their exploratory walking and overall metabolic rate, as a strategy to maximize the net energy intake. This study integrates behavioral and metabolic compensations as a response to foraging at different costs in natural conditions.     

Reversed optimality and predictive ecology: burrowing depth forecasts population change in a bivalve

Optimality reasoning from behavioural ecology can be used as a tool to infer how animals perceive their environment. Using optimality principles in a 'reversed manner' may enable ecologists to predict changes in population size before such changes actually happen. Here we show that a behavioural anti-predation trait (burrowing depth) of the marine bivalve Macoma balthica can be used as an indicator of the change in population size over the year to come. The per capita population growth rate between years t and t+1 correlated strongly with the proportion of individuals living in the dangerous top 4 cm layer of the sediment in year t: the more individuals in the top layer, the steeper the population decline. This is consistent with the prediction based on optimal foraging theory that animals with poor prospects should accept greater risks of predation. This study is among the first to document fitness forecasting in animals.     

Effects of body size and sociality on the anti‐predator behaviour of foraging bees

Pollinators, like most other animals, often face a tradeoff between increasing food uptake and minimising predation. An earlier model suggests that social bees should be more likely than solitary bees to adopt riskier foraging strategies in order to increase food uptake. In this paper, we extend this model by studying the effect of body size, in addition to sociality, on the predation–intake rate tradeoff. When, following standard practice, we express the foraging strategies in terms of mortality probability and net food uptake, we find that body size should have no effect on the foraging strategies of solitary bees. Social bees, on the other hand, should change their foraging preferences according to their size. Small social bees should tend to maximise food uptake, and large social bees to minimise mortality rate. Mortality, however, is the product of two terms: the probability of suffering an attack and the probability of succumbing to it. Noting that larger bees are less susceptible to succumb to a predation attempt than smaller bees, model predictions change when foraging strategies are expressed in terms of exposure to predators. Following this second approach, exposure to predators should increase monotonically with body size in solitary bees. In social bees it should reach a minimum for medium‐sized bees. We conclude that both bee body size and sociality should be considered when studying the effect of predators on resource use.