饥饿和食物单宁酸对东方田鼠(Microtus fortis)食物摄入量和觅食行为的影响

实验室条件下,测定饥饿和食物单宁酸对东方田鼠食物摄入量和觅食行为的影响。结果表明,饥饿使实验个体的食物总摄入量增加,食物摄入率及口量大小随饥饿强度的增大而增加,而觅食频次则无显著改变,实验个体每取食回合的觅食时间呈缓慢增加的趋势,与对照组比较,觅食时间差异不显著。东方田鼠优先选择0％单宁酸食物,次为3%单宁酸食物,而对6%单宁酸食物的摄入量最少。在饥饿条件下,东方田鼠食物摄入率的增加主要源于其口量大小,觅食频次和觅摄食时间对食物摄入量增加的贡献不显著。在饥饿条件下,植食性小哺乳动物并未通过延长觅食时间,降低用于防卫、繁殖活动时间来增加食物摄入量,而是通过增加口量大小,提高其食物摄入率来满足其营养需要。验证了饥饿与植物次生化合物共同作用引起田鼠类动物生理的改变,能影响其食物摄入量及觅食行为的假设。     

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。     

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。     

东方田鼠家族群对成员个体觅食行为的影响

植食性哺乳动物在分享社群觅食带来好处的同时,是否因个体间的相互干扰而影响其摄入率。在新鲜马唐叶片构建的均质密集食物斑块上,测定东方田鼠家族群成员个体在食物斑块上的觅食行为序列过程及行为参数,检验家族群存在对成员个体觅食行为的影响。结果发现,东方田鼠家族群雌、雄成员个体的觅食行为参数均无显著差异。然而,与单只个体相比,家族群觅食尽管能显著地缩短成员个体的觅食决定时间,但却显著地降低了成员个体的摄入率。分析觅食行为参数觅食中断时间发现,相较于单只个体,家族群成员个体间因相互干扰而引起的觅食中断时间的增加,不但增大了收获每口食物的时间,而且导致其摄入率下降。检测家族群成员个体各警觉行为动作参数,发现,成员个体间的相互干扰能引致个体的一般扫视、盯视及嗅闻动作时间比例显著增大,尽管直立扫视和静听动作时间比例减少显著,但并未使个体的觅食中断时间减小。结果充分说明,东方田鼠家族群成员个体间的相互干扰能使个体觅食行为参数发生变异,导致觅食中断时间增加,摄入率降低。     

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。     

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。     

东方田鼠警觉对其功能反应的作用格局

植食性哺乳动物功能反应描述了摄入率与植物可利用性变量的动态关系。动物警觉因占用了处理食物时间即觅食中断时间而延迟与下一口食物相遇,引致摄入率降低,进而对功能反应构型产生影响。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,建立功能反应函数模型,检验觅食中断对功能反应的作用格局及机制。结果发现,除小叶片斑块觅食中断时间在觅食活动中所占的比例较低外,在大、中型叶片斑块的觅食中断时间比例均达到15.42%-26.82%;尽管,觅食中断使摄入率降低了33%,但东方田鼠功能反应仍为Ⅱ型功能反应。除东方田鼠采食时间及觅食回合时间随叶片重增大保持相对稳定外,处理时间及觅食中断时间均随叶片重增大呈线性递增趋势;采食时间、处理时间及觅食中断时间随口量的增大呈线性递增趋势;采食率随叶片重和口量增大呈指数递减趋势。研究结果揭示,东方田鼠因警觉引起的觅食中断事件是导致采食率及摄入率降低的主要因子。摄入率测定值与模型预测值的线性回归极显著(P < 0.01),表明,新建立的功能反应模型具有良好的可预测性。推测,东方田鼠因警觉引起采食率及摄入率减小的代价,是以延长觅食时间来补偿的。研究结果充分验证了,在可利用性植物密集斑块,由植物大小调控的动物口量决定其摄入率,且受采食和处理食物竞争及觅食中断的制约,其功能反应为Ⅱ型功能反应的假说。     

单宁酸对根田鼠食物摄入量和蛋白质消化率的效应

在食物中含10 %和20%蛋白质的条件下, 采用食物平衡法测定了单宁酸对根田鼠食物摄入量和蛋白质消化率的作用。食物蛋白质含量为10 %时, 第1～5 天, 单宁酸对根田鼠食物摄入量具有显著的抑制作用, 自第6天, 其作用不明显, 以3 %和6 %单宁酸处理的根田鼠, 其食物蛋白质消化率较对照组分别降低22 %和47.67 %;在食物蛋白质含量为20 %的条件下, 单宁酸对根田鼠的食物摄入量和蛋白质消化率无显著作用。上述结果验证了植物次生化合物能抑制植食性小哺乳动物食物摄入量及蛋白质消化率的假设。     

在不同营养状态下东方田鼠对陌生食物选择学习的研究

通过不同饥饿处理 ,研究了东方田鼠在不同营养状态下 ,对陌生食物尝试和选择的行为模式。结果表明 ,初次遇到陌生食物时 ,各处理组对陌生食物的采食量分别为饱食 6.9± 1 .8g,饥饿组为 4.8± 1 .6g,饥饿 食物短缺组为 3.4± 1 .4g。各处理组动物对陌生食物的摄入量间差异极为显著 ( P=0 .0 0 4 3)。饱食组、饥饿组及饥饿 食物短缺组动物初次遇到陌生食物时 ,所摄入的陌生食物量占总摄入量的百分比分别为72 %、38%与 1 2 %。说明 ,在实验室条件下饱食动物较饥饿和饥饿 食物短缺的动物学习选择摄食陌生食物的倾向较强。     

食物单宁酸对根田鼠(Microtus oeconomus)钠平衡的作用

验证了单宁酸能引起植食性哺乳动物钠平衡的假设.在两种食物蛋白质水平条件下,测定了摄食含0%,3%和6%单宁酸食物的根田鼠的钠平衡和肾上腺肾小球的大小.结果表明,食物单宁酸能显著影响根田鼠的钠平衡,钠的丢失主要通过尿液.同时,单宁酸能显著影响试验个体的肾上腺肾小球的体积.初步研究显示,植食性哺乳动物对摄入的植物次生化合物的解毒引起机体钠平衡失调,而非植物次生化合物的消化抑制作用所致.     

Sward height and bite size affect the functional response of barnacle geese Branta leucopsis

Intake rate, the rate in which herbivores can process their food, is presumed to be an important factor in habitat selection down to the scale of the foraging patch. Much attention has been given to the selection of swards of high nutritional quality, but much less has been given to the influences of sward structure on patch selection in small herbivores. In this study we tested the effects of sward density and height on the functional foraging response of barnacle geese, Branta leucopsis. The functional response curve for herbivores describes how intake rate is affected by food availability. We conducted feeding trials to determine intake rate and bite size of barnacle geese on experimentally manipulated swards. Results indicate that intake rate is mainly dependent on sward height and that there is a strong correlation between bite size and intake rate. Sward density does not influence the rate of food consumption; it is, however, a crucial parameter affecting potential total yield. We conclude that bite size is the crucial parameter influencing intake rate. Bite size is explained both by sward height and individual differences in bill morphology. Furthermore, intake rate seems to be dependent on the physical structure of the grass species consumed.     

实验检验东方田鼠学习能力的性别差异

东方田鼠选择从禾本科植物叶片的不同部位采食会引起食物摄入率的巨大差异,这项具有明显的认知和学习行为的觅食活动是否存在性别差异。在由带两片叶的新鲜马唐茎秆配置的密集食物斑块上,测定东方田鼠觅食马唐叶片的行为序列、过程及参数,检验其在学习能力上的性别差异。结果表明,雌鼠和雄鼠的觅食决定时间尽管均随觅食学习天数的增加而减小,但雌鼠的觅食决定时间显著地高于雄鼠的;雌鼠和雄鼠均由从对叶片部位无差别选择和采食转向偏爱从叶片基部采食。学会这项采食技能后,雌鼠从叶片基部采食的选择指数显著地高于雄鼠的;二者的摄入率均随觅食学习天数的增加极显著地增大,但雌鼠的摄入率却极显著地高于雄鼠的。研究结果充分验证了提出的假设:雄鼠较雌鼠能更快地习得从禾本科植物叶片的基部采食和收获食物;但当二者习得这项采食技艺后,雌鼠较雄鼠能获得更高的食物摄入率;同时也说明,新建立的评价小型植食性哺乳动物认知和学习能力的实验方法,具有可靠的有效性和实用性。     

Sideroxylonal in Eucalyptus foliage influences foraging behaviour of an arboreal folivore

Plant secondary metabolites (PSMs) offer plants chemical defences against herbivores, and are known to influence intake and diet choice in both insect and mammalian herbivores. However, there is limited knowledge regarding how PSMs influence herbivore foraging decisions. Herbivore foraging decisions, in turn, directly impact on which individual plants, and plant species, are selected for consumption. We took advantage of the natural variation in sideroxylonal concentrations in the foliage of Eucalyptus melliodora (Cunn. ex Schauer) to investigate feeding patterns of a marsupial folivore, the common ringtail possum, Pseudocheirus peregrinus (Boddaert 1785). Foliage, collected from six trees, contained between 0.32 and 12.97 mg g-DM^-1 sideroxylonal. With increasing sideroxylonal concentrations, possums decreased their total intake, rate of intake and intake per feeding bout, and increased their cumulative time spent feeding. Possums did not alter their total feeding time, number of feeding bouts or time per feeding bout in response to increasing sideroxylonal concentrations. Results demonstrate important behavioural changes in foraging patterns in response to sideroxylonal. These behavioural changes have important implications, in relation to altered foraging efficiency and potential predation risk, for herbivores foraging in the field. As a result, the spatial distribution of dietary PSMs across a landscape may directly influence herbivore fitness, and ultimately habitat selection of mammalian herbivores.     

Moose and vole browsing patterns in experimentally assembled pure and mixed forest stands

Plants growing in diverse communities are believed to exhibit associational resistance to herbivores, but this hypothesis has seldom been tested experimentally for vertebrate herbivores in forest ecosystems. We examined browsing patterns of the two principal mammalian herbivores of Finnish boreal forests, moose and voles, in young stands where tree species diversity and composition were experimentally manipulated. The stands were composed either of monocultures or different 2–5 species mixtures of Norway spruce, Scots pine, Siberian larch, silver birch, and black alder. Voles and moose showed contrasting responses to stand diversity and species composition. In accordance with the predictions of the associational resistance hypothesis, vole damage was higher in tree monocultures than in mixed stands, although stand diversity effects were statistically significant only at one of the three study areas. Voles also damaged more trees in coniferous than in deciduous stands. In contrast, moose browsing tended to increase with the number of tree species in a stand and with the presence of the preferred tree species, birch, in a mixture. The observed differences in vole and moose responses to stand diversity and species composition are likely to be due to different feeding specialisation, foraging patterns, and movement ability of these herbivores. Voles switched to trees only when the supply of a more preferred food (grasses and forbs) was depleted and restricted their feeding choice only to the most palatable tree species regardless of the number of tree species present per stand. In contrast, tree branches and foliage represented an important part of moose diet throughout the year; moose may be able to tolerate secondary plant metabolites of different tree species better than voles and may thus benefit from diet broadening when more tree species are available. Furthermore, the home range size and foraging ability of these two very differently sized herbivores may partly explain the observed differences in utilisation of different tree species. Finally, both moose and voles showed high spatial and temporal variation in their feeding; in particular, vole damage was more influenced by tree species diversity in areas and years with high vole densities. Thus, diversification of forest stands may have very different effects on mammalian browsing depending on the herbivores present, their densities, and the tree species used in reforestation.     

Experimental demonstration of the antiherbivore effects of silica in grasses: impacts on foliage digestibility and vole growth rates

The impact of plant-based factors on the population dynamics of mammalian herbivores has been the subject of much debate in ecology, but the role of antiherbivore defences in grasses has received relatively little attention. Silica has been proposed as the primary defence in grasses and is thought to lead to increased abrasiveness of foliage so deterring feeding, as well as reducing foliage digestibility and herbivore performance. However, at present there is little direct experimental evidence to support these ideas. In this study, we tested the effects of manipulating silica levels on the abrasiveness of grasses and on the feeding preference and growth performance of field voles, specialist grass-feeding herbivores. Elevated silica levels did increase the abrasiveness of grasses and deterred feeding by voles. We also demonstrated, for the first time, that silica reduced the growth rates of both juvenile and mature female voles by reducing the nitrogen they could absorb from the foliage. Furthermore, we found that vole feeding leads to increased levels of silica in leaves, suggesting a dynamic feedback between grasses and their herbivores. We propose that silica induction due to vole grazing reduces vole performance and hence could contribute to cyclic dynamics in vole populations.     

植物化学防卫与植食性哺乳动物的适应对策

植物化学防卫与植食性哺乳动物的适应是动物—植物系统协同进化研究的重要内容。植物次生化合物可降低动物的食物摄入量及消化率、蛋白质可利用率。某些次生化合物还影响植食性哺乳动物的正常繁殖活动。单宁是动物的重要觅食阻遏剂之一。动物在学习食物选择的过程中,通过认知过程和感知过程处理食物信息,选择食物项目。幼体在胚胎期和哺乳期能从母体获得食物信息,或向有觅食经验的同胞伙伴学习处理食物的经验。动物亦可通过形成络和物,改变体内环境,通过微生物降解、氧化还原、基础代谢率等降低生理对策,以降低植物次生化和物的影响。     

Importance of nutritional and anti-parasite strategies in the foraging decisions of horses: an experimental test

The primary goal of foraging herbivores is to maximise the net rate of intake of digestible energy (or of a limiting nutrient). However, foraging strategies of herbivores are also sensitive to other selective forces (e.g. predation, parasites), which may modify their choice of feeding patches. Horses feed in spatially complex swards, and allocate their time among patches which vary both in terms of their nutritional benefits, and the risk of parasitism. It has long been suggested that horses allocate time among patches principally in relation to the risk of parasitism, though the nutritional costs and benefits of different foraging choices must play some role too. In this study, we investigated the roles of nutritional and anti-parasite factors in foraging decisions by horses. Six naturally parasitized and six unparasitized two-year-old geldings were allowed to graze from pairs of trays (112×72 cm) with swards at two heights (tall: 52 cm, medium: 15 cm) and two levels of contamination (280 g m⁻² of faeces, no faeces) in a latin square, pair-wise design with six different choices. In nutritional terms the differences between the swards were slight: the tall sward provided larger bite sizes and higher intake rates (+36%). Feeding motivation was not high, and the faecal stimulus was strong. The horses selected uncontaminated swards when the nutritional benefits were identical, and tall swards in virtually all circumstances. Contamination of the preferred tall sward (i.e. a test of the tradeoff between anti-parasite and nutritional strategies) had little effect on its selection by the horses. Their parasite status also had little effect: both groups selected the tall sward in the tradeoff (though this was only a trend for the non-parasitised group in one analysis). We conclude that patch selection by horses in these particular conditions was driven principally by their nutritional strategy.     

Foraging costs of vigilance in large mammalian herbivores

Vigilance has been assumed to reduce food intake by taking away time from food processing. Such foraging costs of vigilance have been predicted to have profound effects on the structure of communities. Recently, however, it has been argued that mammalian herbivores might be capable of maintaining their rate of food intake despite being vigilant, because of their ability to scan the environment while chewing vegetation. We conducted behavioral observations to evaluate whether vigilance decreases the bite rate of free-ranging female bison ( Bison bison ) in Prince Albert National Park and elk ( Cervus canadensis ) in Yellowstone National Park. Modeling of foraging processes indicated that chewing time exceeded the time that bison and elk spent searching for food, interacting with conspecifics, and scanning. Consequently, bison and elk might have been capable of maintaining their rate of food intake despite vigilance. The maintenance of intake rate would have required bison and elk to match scanning events closely with chewing bouts, but we did not detect a positive correlation between the duration of scanning bouts and the number of consecutive bites taken just before vigilance events. As a result, vigilance was costly, and as it increased, bite rate declined for both herbivore species. Scanning still overlapped partially with food handling. Indeed, we estimated that 31% of feeding time being vigilant decreased bite rate by 20% for bison and 26% for elk, whereas total absence of overlap between chewing and scanning should have reduced bite rate by 31%. While we observed that vigilance induced foraging costs, these costs were less important than traditionally assumed.     

The influence of bite size on foraging at larger spatial and temporal scales by mammalian herbivores

Organisms respond to their heterogeneous environment in complex ways at many temporal and spatial scales. Here, I examine how the smallest scale process in foraging by mammalian herbivores, taking a bite, influences plants and herbivores over larger scales. First, because cropping bites competes with chewing them, bite size influences short-term intake rate of herbivores within plant patches. On the other hand, herbivores can chew bites while searching for new ones, thus influencing the time spent vigilant and intake rate as animals move among food patches. Therefore, bite size affects how much time herbivores must spend foraging each day. Because acquiring energy is necessary for fitness, herbivores recognize the importance of bite size and select bites, patches and diets based on tradeoffs between harvesting rates, digestion, and sheering forces. In turn, induced structural defenses of plants, such as thorns, allow plants to respond immediately to herbivory by reducing bite size and thus tissue loss. Over evolutionary time, herbivores have adapted mouth morphology that allows them to maximize bite size on their primary forage plant, whereas plants faced with large mammalian herbivores have adapted structures such as divarication that minimize bite size and protect themselves from herbivory. Finally, bite size available among plant communities can drive habitat segregation and migration of larger herbivores across landscapes.