Flight metabolic rate in the Glanville fritillary butterfly

Dispersal is a highly important life history trait. In fragmented landscapes the long-term persistence of populations depends on dispersal. Evolution of dispersal is affected by costs and benefits and these may differ between different landscapes. This results in differences in the strength and direction of natural selection on dispersal in fragmented landscapes. Dispersal has been shown to be a nonrandom process that is associated with traits such as flight ability in insects. This thesis examines genetic and physiological traits affecting dispersal in the Glanville fritillary butterfly (Melitaea cinxia). Flight metabolic rate is a repeatable trait representing flight ability. Unlike in many vertebrates, resting metabolic rate cannot be used as a surrogate of maximum metabolic rate as no strong correlation between the two was found in the Glanville fritillary. Resting and flight metabolic rate are affected by environmental variables, most notably temperature. However, only flight metabolic rate has a strong genetic component. Molecular variation in the much-studied candidate locus phosphoglucose isomerase (Pgi), which encodes the glycolytic enzyme PGI, has an effect on carbohydrate metabolism in flight. This effect is temperature dependent: in low to moderate temperatures individuals with the heterozygous genotype at the single nucleotide polymorphism (SNP) AA111 have higher flight metabolic rate than the common homozygous genotype. At high temperatures the situation is reversed. This finding suggests that variation in enzyme properties is indeed translated to organismal performance. High-resolution data on individual female Glanville fritillaries moving freely in the field were recorded using harmonic radar. There was a strong positive correlation between flight metabolic rate and dispersal rate. Flight metabolic rate explained one third of the observed variation in the one-hour movement distance. A fine-scaled analysis of mobility showed that mobility peaked at intermediate ambient temperatures but the two common Pgi genotypes differed in their reaction norms to temperature. As with flight metabolic rate, heterozygotes at SNP AA111 were the most active genotype in low to moderate temperatures. The results show that molecular variation is associated with variation in dispersal rate through the link of flight physiology under the influence of environmental conditions. The evolutionary pressures for dispersal differ between males and females. The effect of flight metabolic rate on dispersal was examined in both sexes in field and laboratory conditions. The relationship between flight metabolic rate and dispersal rate in the field and flight duration in the laboratory were found to differ between the two sexes. In females the relationship was positive, but in males the longest distances and flight durations were recorded for individuals with low flight metabolic rate. These findings may reflect male investment in mate locating. Instead of dispersing, males with high flight metabolic rate may establish territories and follow a perching strategy when locating females and hence move less on the landscape level. Males with low metabolic rate may be forced to disperse due to low competitive success or may show adaptations to an alternative strategy: patrolling. In the light of life history trade-offs and the rate of living theory having high metabolic rate may carry a cost in the form of shortened lifespan. Experiments relating flight metabolic rate to longevity showed a clear correlation in the opposite direction: high flight metabolic rate was associated with long lifespan. This suggests that individuals with high metabolic rate do not pay an extra physiological cost for their high flight capacity, rather there are positive correlations between different measures of fitness. These results highlight the importance of condition.

Changes in catch rates and length and age at maturity, but not growth, of an estuarine plotosid (Cnidoglanis macrocephalus) after heavy fishing

The hypothesis that heavy fishing pressure has led to changes in the biological characteristics of the estuary cobbler (Cnidoglanis macrocephalus) was tested in a large seasonally open estuary in southwestern Australia, where this species completes its life cycle and is the most valuable commercial fish species. Comparisons were made between seasonal data collected for this plotosid (eeltail catfish) in Wilson Inlet during 2005–08 and those recorded with the same fishery-independent sampling regime during 1987–89. These comparisons show that the proportions of larger and older individuals and the catch rates in the more recent period were far lower, i.e., they constituted reductions of 40% for fish ≥430 mm total length, 62% for fish ≥4 years of age, and 80% for catch rate. In addition, total mortality and fishing-induced mortality estimates increased by factors of ~2 and 2.5, respectively. The indications that the abundance and proportion of older C. macrocephalus declined between the two periods are consistent with the perception of long-term commercial fishermen and their shift toward using a smaller maximum gill net mesh to target this species. The sustained heavy fishing pressure on C. macrocephalus between 1987–89 and 2005–08 was accompanied by a marked reduction in length and age at maturity of this species. The shift in probabilistic maturation reaction norms toward smaller fish in 2005–08 and the lack of a conspicuous change in growth between the two periods indicate that the maturity changes were related to fishery-induced evolution rather than to compensatory responses to reduced fish densities.

异质性环境中两种匍匐茎草本植物的克隆可塑性

　　克隆植物常常生长在资源异质性分布的环境中，克隆植物特有的生活史特征使其对环境和资源变化产生的反应规范（Reaction norm）具有特殊的表现和适应意义。本文以匍匐茎草本植物蛇莓(Duchesnea indica)和活血丹(Glechoma longituba)为材料，研究克隆植物在不同异质性环境中的反应规范，即可塑性（plasticity）和克隆分株间相互作用（即克隆整合（clonal integration））对表型的影响及其适应意义。 蛇莓和活血丹是具有不同分枝类型的两种匍匐茎草本植物。对于匍匐茎草本植物而言，生境中光资源分布在时间和空间上的变化极为丰富。本文主要研究在不同环境中，这两种克隆植物克隆分株间的相互作用是如何通过影响其形态特征，从而影响其光资源获取策略。 实验涉及了4种光照条件，分为5个部分，分别在大棚和野外进行。 1．同质光环境中蛇莓和活血丹克隆片段的可塑性 在大棚中，将整个克隆片段（clonal fragment）都放置在光照强度分别为自然光照PPFD（Photosynthetic Photon Flux Density）的100％、50％和10％三种不同的同质光环境中，目标是通过比较不同光照强度同质环境中不同基因型的形态特征，揭示匍匐茎草本克隆植物的反应规范及遗传差异。实验结果发现，当光照强度从100% PPFD 降到 50% PPFD时，活血丹增加对地上部分生物量投资，增加部分多投入到匍匐茎上，用于支持匍匐茎实现觅食行为，叶片和叶柄的变化不大。同样条件下，蛇莓的生物量投资却向根部转移，地上部分叶片和叶柄形态上的变化比匍匐茎的形态变化显著，出现增强对光资源吸收能力的变化。当光照强度从50%降到 10%时，活血丹继续增加对地上部分的生物量投资，但由于光照强度弱，几乎不足以维持新的匍匐茎的发生，所有增加的生物量部分都投入到叶片和叶柄的变化上。这种条件下，蛇莓的生物量投资依然向根部转移，叶片和叶柄为增加对光资源吸收能力而发生了相应变化，匍匐茎的新增数减少，形态变化也不大。实验证明，可能由于匍匐茎的发生方式不同，活血丹的匍匐茎节间长度，叶片和叶柄的关联程度比蛇莓相应性状的关联程度强。不同基因型的蛇莓和活血丹对光照梯度的反应有显著差异。 2．异质光环境中蛇莓克隆片段的可塑性 针对克隆植物分株间在相当长的时间里形体相连，而环境异质性可能存在于分株大小的空间尺度，设置异质环境，将蛇莓的不同相连分株放置在不同的光资源斑块中，研究其整合作用在适应异质性环境过程中的作用及其对表型的影响。首先设置同一种资源（光资源）的梯度差异斑块。 在光斑块中，整合作用发生与否、作用方式与资源梯度、分株自身所处斑块的资源条件和分株的年龄结构有关。在不同的条件下，整合作用可能造成对表型的不同影响，从而可能加剧或削弱分株对本地资源的反应（局部反应，local response）。整合作用对分株局部反应的影响强度和方向，在不同基因型间存在明显差异。这些基因型间的差异暗示，整合作用可能也是具有遗传基础的一种独立性状。 3．光块斑和养分斑块共存环境中活血丹克隆片段的可塑性 设置光和养分的资源互补性斑块。将相连分株种植在不同斑块中。当生长在高光低养斑块中的分株与其互补斑块（低光高养斑块）的分株相连时，其适合度相关性状的值增加，根冠比是可塑的，通过相应的形态变化，高光照斑块中的分株捕获光资源的能力增强，但两种斑块中植株的吸收养分的能力却没有大的变化。收益－损耗分析显示整合作用有益于异质环境中的植株，低光高养斑块中植株的生物量获益。实验结果证明了异质环境中相连分株间存在光合产物和养分传递。在这种环境种，分株形态的变化对本地斑块发生趋富反应，形态的改变有助于对本地丰富资源的吸收。这种反应有利于克隆片段对资源的吸收。 4．野外环境中蛇莓克隆片段的可塑性 基于大棚实验资料，对野外林下蛇莓种群进行监测。首先对单株在林下复杂的光环境中的表现进行了跟踪，结果发现，随着分株数目的增加，适合度相关性状值有所提高，而处于不同位置的相连分株在形态上的差异不显著。大棚实验中发现的显著形态变化在具有显著差异的自然环境条件下没有发生，这可能暗示着在资源条件变化频率较高的环境中整合作用的作用方式。分株不是对某一时刻资源条件进行形态特化，而是通过对所扩展的总体区域的总的环境条件调整表型。这样，从克隆片段的水平上看，资源的吸收可能达到较高水平。在密度增大，植株间互相荫蔽使得光照减弱的情况下，匍匐茎的变化没有表现出觅食行为，叶片和叶柄出现增大增长等增强吸收光能力的相应表型变化，证明叶片叶柄是克服光照不足的主要器官。这些结果与大棚实验结果一致。蛇莓基株对于相连分株内部可能具有密度调节功能，从而通过减弱叶片增大的趋势，增长匍匐茎，分散新生分株，使密度保持在一定程度，不对种群的发展造成阻碍作用。非相连分株间随着密度的增大，相互间的作用方式类似于非克隆植物：出现叶片增大，叶柄增长的现象，同时密度制约了适合度相关性状如分株数目等的增加。 5．野外环境中蛇莓种群格局动态 最后，在进化的单位，种群水平上对蛇莓种群在自然界的动态进行了调查。Spearman相关分析，没有发现光资源和测量指标在量上的相关。在自然界中整合作用使大量分散分布、相互连接的分株相互作用，以促进基株对资源的获取。Moran’s I指数分析显示，随时间的推进，各个指标在相关尺度上有所增加。这暗示了整合作用的存在，分株间联系的加强。大棚实验中观察到的关于各个器官的作用的结论在自然界中得到进一步的验证。蛇莓匍匐茎在遮荫环境中不是作为觅食器官。为了争取对更多光资源的吸收，蛇莓在叶片数和叶形态上发生改变。对分株数、叶片数、匍匐茎数目的分布格局的调查显示出没有一定的规律性，而且处于不断的变化中。这一结果可能暗示种群发展过程中处理异质性分布资源的对策的变化。 此外，本文还揭示了可塑性和整合作用在基因型间的差异，针对所发现的现象，从作用的遗传机制上对可塑性及其适应意义进行了讨论。结合分子生物学和地统计学等学科的研究成果，对进一步的实验提出了方法和路线。

How does developmental plasticity differ between diverse types of nutrition regimes?

Tese de mestrado em Biologia Evolutiva e do Desenvolvimento, apresentada à Universidade de Lisboa, através da Faculdade de Ciências, 2016

MHC class I expression dependent on bacterial infection and parental factors in whitefish embryos (Salmonidae).

Ecological conditions can influence not only the expression of a phenotype, but also the heritability of a trait. As such, heritable variation for a trait needs to be studied across environments. We have investigated how pathogen challenge affects the expression of MHC genes in embryos of the lake whitefish Coregonus palaea. In order to experimentally separate paternal (i.e. genetic) from maternal and environmental effects, and determine whether and how stress affects the heritable variation for MHC expression, embryos were produced in full-factorial in vitro fertilizations, reared singly, and exposed at 208 degree days (late-eyed stage) to either one of two strains of Pseudomonas fluorescens that differ in their virulence characteristics (one increased mortality, while both delayed hatching time). Gene expression was assessed 48 h postinoculation, and virulence effects of the bacterial infection were monitored until hatching. We found no evidence of MHC class II expression at this stage of development. MHC class I expression was markedly down-regulated in reaction to both pseudomonads. While MHC expression could not be linked to embryo survival, the less the gene was expressed, the earlier the embryos hatched within each treatment group, possibly due to trade-offs between immune function and developmental rate or further factors that affect both hatching timing and MHC expression. We found significant additive genetic variance for MHC class I expression in some treatments. That is, changes in pathogen pressures could induce rapid evolution in MHC class I expression. However, we found no additive genetic variance in reaction norms in our study population.

Developmental, morphological, and behavioural plasticity in the reproductive strategies of stink bugs and their egg parasitoids

L’environnement façonne la physiologie, la morphologie et le comportement des organismes par l’entremise de processus écologiques et évolutifs complexes et multidimensionnels. Le succès reproducteur des animaux est déterminé par la valeur adaptative d’un phénotype dans un environnement en modification constante selon une échelle temporelle d’une à plusieurs générations. De plus, les phénotypes sont façonnés par l’environnement, ce qui entraine des modifications adaptatives des stratégies de reproduction tout en imposant des contraintes. Dans cette thèse, considérant des punaises et leurs parasitoïdes comme organismes modèles, j’ai investigué comment plusieurs types de plasticité peuvent interagir pour influencer la valeur adaptative, et comment la plasticité des stratégies de reproduction répond à plusieurs composantes des changements environnementaux (qualité de l’hôte, radiation ultraviolette, température, invasion biologique). Premièrement, j’ai comparé la réponse comportementale et de traits d’histoire de vie à la variation de taille corporelle chez le parasitoïde Telenomus podisi Ashmead (Hymenoptera : Platygastridae), démontrant que les normes de réaction des comportements étaient plus souvent positives que celles des traits d’histoires de vie. Ensuite, j’ai démontré que la punaise prédatrice Podisus maculiventris Say (Hemiptera : Pentatomidae) peut contrôler la couleur de ses œufs, et que la pigmentation des œufs protège les embryons du rayonnement ultraviolet; une composante d’une stratégie complexe de ponte qui a évoluée en réponse à une multitude de facteurs environnementaux. Puis, j’ai testé comment le stress thermique affectait la dynamique de la mémoire du parasitoïde Trissolcus basalis (Wollaston) (Hymenoptera : Platygastridae) lors de l’apprentissage de la fiabilité des traces chimiques laissées par son hôte. Ces expériences ont révélé que des températures hautes et basses prévenaient l’oubli, affectant ainsi l’allocation du temps passé par les parasitoïdes dans des agrégats d’hôtes contenant des traces chimiques. J’ai aussi développé un cadre théorique général pour classifier les effets de la température sur l’ensemble des aspects comportementaux des ectothermes, distinguant les contraintes des adaptations. Finalement, j’ai testé l’habileté d’un parasitoïde indigène (T. podisi) à exploiter les œufs d’un nouveau ravageur invasif en agriculture, Halyomorpha halys Stål (Hemiptera : Pentatomidae). Les résultats ont montré que T. podisi attaque les œufs de H. halys, mais qu’il ne peut s’y développer, indiquant que le ravageur invasif s’avère un « piège évolutif » pour ce parasitoïde. Cela pourrait indirectement bénéficier aux espèces indigènes de punaises en agissant comme un puits écologique de ressources (œufs) et de temps pour le parasitoïde. Ces résultats ont des implications importantes sur la réponse des insectes, incluant ceux impliqués dans les programmes de lutte biologique, face aux changements environnementaux.

Changes in catch rates and length and age at maturity, but not growth, of an estuarine plotosid (cnidoglanis macrocephalus) after heavy fishing

The hypothesis that heavy fishing pressure has led to changes in the biological characteristics of the estuary cobbler (Cnidoglanis macrocephalus) was tested in a large seasonally open estuary in southwestern Australia, where this species completes its life cycle and is the most valuable commercial fish species. Comparisons were made between seasonal data collected for this plotosid (eeltail catfish) in Wilson Inlet during 2005-08 and those recorded with the same fishery-independent sampling regime during 1987-89. These comparisons show that the proportions of larger and older individuals and the catch rates in the more recent period were far lower, i.e., they constituted reductions of 40% for fish ≥430 mm total length, 62% for fish ≥4 years of age, and 80% for catch rate. In addition, total mortality and fishing-induced mortality estimates increased by factors of ~2 and 2.5, respectively. The indications that the abundance and proportion of older C. macrocephalus declined between the two periods are consistent with the perception of long-term commercial fishermen and their shift toward using a smaller maximum gill net mesh to target this species. The sustained heavy fishing pressure on C. macrocephalus between 1987-89 and 2005-08 was accompanied by a marked reduction in length and age at maturity of this species. The shift in probabilistic maturation reaction norms toward smaller fish in 2005-08 and the lack of a conspicuous change in growth between the two periods indicate that the maturity changes were related to fishery-induced evolution rather than to compensatory responses to reduced fish densities.

Phenological response of sea turtles to environmental variation across a species' northern range

Variations in environmental parameters (e. g. temperature) that form part of global climate change have been associated with shifts in the timing of seasonal events for a broad range of organisms. Most studies evaluating such phenological shifts of individual taxa have focused on a limited number of locations, making it difficult to assess how such shifts vary regionally across a species range. Here, by using 1445 records of the date of first nesting for loggerhead sea turtles (Caretta caretta) at different breeding sites, on different continents and in different years across a broad latitudinal range (25-39 degrees ' N), we demonstrate that the gradient of the relationship between temperature and the date of first breeding is steeper at higher latitudes, i.e. the phenological responses to temperature appear strongest at the poleward range limit. These findings support the hypothesis that biological changes in response to climate change will be most acute at the poleward range limits and are in accordance with the predictions of MacArthur's hypothesis that poleward range limit for species range is environmentally limited. Our findings imply that the poleward populations of loggerheads are more sensitive to climate variations and thus they might display the impacts of climate change sooner and more prominently.

Personality and plasticity: consistent responses within-, but not across-temperature situations in crabs

Recent research suggests that repeated assays of behaviour, conducted both within and across situations, could reveal important insights into two traditionally distinct areas of study: animal personality and individual differences in behavioural plasticity. However, at present relatively few studies present such data, and few consider how changing abiotic conditions affect behavioural plasticity. Individual differences in metabolic rate have been suggested as a proximate mechanism promoting personality, leading one to speculate that individual differences in metabolic sensitivity to temperature may affect behavioural responses in ectotherms. At present, only one study (out of two) has tested for and shown individual differences in behavioural responses to temperature. Here, we repeatedly assayed the behaviour of a marine crab across a narrow range of temperatures to test for individual differences in responses to temperature. We observed large inter-individual differences in behaviour that were consistent over time at a given temperature (evidence for personality), and individual differences in responses to temperature (evidence for plasticity). This study adds to the very scant literature on ectotherm behavioural sensitivity to temperature, and suggests the phenomenon might be widespread. We speculate about the role of metabolism as a proximate mechanism that might explain these individual differences in plasticity and make suggestions for future research to test this hypothesis. © Koninklijke Brill NV, Leiden, The Netherlands.

Individual variation in thermal performance curves: swimming burst speed and jumping endurance in wild-caught tropical clawed frogs

The importance of studying individual variation in locomotor performance has long been recognized as it may determine the ability of an organism to escape from predators, catch prey or disperse. In ectotherms, locomotor performance is highly influenced by ambient temperature (Ta), yet several studies have showed that individual differences are usually retained across a Ta gradient. Less is known, however, about individual differences in thermal sensitivity of performance, despite the fact that it could represent adaptive sources of phenotypic variation and/or additional substrate for selection to act upon. We quantified swimming and jumping performance in 18 wild-caught tropical clawed frogs (Xenopus tropicalis) across a Ta gradient. Maximum swimming velocity and acceleration were not repeatable and individuals did not differ in how their swimming performance varied across Ta. By contrast, time and distance jumped until exhaustion were repeatable across the Ta gradient, indicating that individuals that perform best at a given Ta also perform best at another Ta. Moreover, thermal sensitivity of jumping endurance significantly differed among individuals, with individuals of high performance at low Ta displaying the highest sensitivity to Ta. Individual differences in terrestrial performance increased with decreasing Ta, which is opposite to results obtained in lizards at the inter-specific and among-individual levels. To verify the generality of these patterns, we need more studies on individual variation in thermal reaction norms for locomotor performance in lizards and frogs.

Using repeatability to study physiological and behavioural traits: ignore time-related change at your peril

© 2015 The Association for the Study of Animal Behaviour. Broad sense repeatability, which refers to the extent to which individual differences in trait scores are maintained over time, is of increasing interest to researchers studying behavioural or physiological traits. Broad sense repeatability is most often inferred from the statistic R (the intraclass correlation, or narrow sense repeatability). However, R ignores change over time, despite the inherent longitudinal nature of the data (repeated measures over time). Here, we begin by showing that most studies ignore time-related change when estimating broad sense repeatability, and estimate R with low statistical power. Given this problem, we (1) outline how and why ignoring time-related change in scores (that occurs for whatever reason) can seriously affect estimates of the broad sense repeatability of behavioural or physiological traits, (2) discuss conditions in which various indices of R can or cannot provide reliable estimates of broad sense repeatability, and (3) provide suggestions for experimental designs for future studies. Finally, given that we already have abundant evidence that many labile traits are 'repeatable' in that broad sense (i.e. R>. 0), we suggest a shift in focus towards obtaining robust estimates of the repeatability of behavioural and physiological traits. Given how labile these traits are, this will require greater experimental (and/or statistical) control and larger sample sizes in order to detect and quantify change over time (if present).

Interação genótipo ambiente para perímetro escrotal em bovinos da raça Nelore utilizando normas de reação

Pós-graduação em Zootecnia - FCAV

Estudo da interação genótipo-ambiente por meio de modelos de normas de reação em bovinos de corte da raça Nelore

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Interação genótipo ambiente sobre características reprodutivas de fêmeas da raça Nelore

Pós-graduação em Genética e Melhoramento Animal - FCAV

Interação genótipo-ambiente para peso ao sobreano na raça Canchim

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)