Co‐evolution of cerebral and cerebellar expansion in cetaceans

Cetaceans possess brains that rank among the largest to have ever evolved, either in terms of absolute mass or relative to body size. Cetaceans have evolved these huge brains under relatively unique environmental conditions, making them a fascinating case study to investigate the constraints and selection pressures that shape how brains evolve. Indeed, cetaceans have some unusual neuroanatomical features, including a thin but highly folded cerebrum with low cortical neuron density, as well as many structural adaptations associated with acoustic communication. Previous reports also suggest that at least some cetaceans have an expanded cerebellum, a brain structure with wide‐ranging functions in adaptive filtering of sensory information, the control of motor actions, and cognition. Here, we report that, relative to the size of the rest of the brain, both the cerebrum and cerebellum are dramatically enlarged in cetaceans and show evidence of co‐evolution, a pattern of brain evolution that is convergent with primates. However, we also highlight several branches where cortico‐cerebellar co‐evolution may be partially decoupled, suggesting these structures can respond to independent selection pressures. Across cetaceans, we find no evidence of a simple linear relationship between either cerebrum and cerebellum size and the complexity of social ecology or acoustic communication, but do find evidence that their expansion may be associated with dietary breadth. In addition, our results suggest that major increases in both cerebrum and cerebellum size occurred early in cetacean evolution, prior to the origin of the major extant clades, and predate the evolution of echolocation.     

The Evolution of Olfactory Capabilities in Wild Birds: A Comparative Study

Modern birds possess an olfactory apparatus similar to that of other vertebrates, yet the major evolutionary forces that drove the evolution of diversity in olfactory capabilities in birds remain elusive. Several non-mutually exclusive evolutionary scenarios for the evolution of olfactory capability in birds have been proposed. Olfactory capability may have evolved due to its role in recognition in social contexts. In addition, olfactory capability may have evolved because it provides a selective advantage in foraging or navigation. Finally, olfactory capability could be favored whenever the ecological conditions of species hindered the use of other senses and/or favored olfaction. Here we evaluate predictions from these hypotheses in a comparative study using a species-level phylogeny of 142 bird species and considering the indirect effects of some predictors on others. We find an interactive effect of aquatic dependence and diet on the size of the olfactory bulb: vegetarian and omnivore birds living in aquatic environments have larger olfactory bulbs than terrestrial birds, whereas species with an animal diet, aquatic and terrestrial species have similar-sized olfactory bulbs. In addition, the size of the olfactory bulb was weakly related with social complexity, with colonial species having relatively larger olfactory bulb than solitary breeders or those forming small aggregations. Our results suggest that the role of foraging in driving enhancements in the avian olfactory apparatus is contingent on ecological conditions that may affect the transmission of odor-based signals. This provides evidence for the largely neglected possibility that the evolution of the olfactory apparatus of birds has been driven by the interaction between ecological and behavioral traits, rather than being solely due to main effects of these traits.     

FORELIMB POSTURE IN DINOSAURS AND THE EVOLUTION OF THE AVIAN FLAPPING FLIGHT-STROKE

Ontogenetic and behavioral studies using birds currently do not document the early evolution of flight because birds (including juveniles) used in such studies employ forelimb oscillation frequencies over 10 Hz, forelimb stroke-angles in excess of 130°, and possess uniquely avian flight musculatures. Living birds are an advanced morphological stage in the development of flapping flight. To gain insight into the early stages of flight evolution (i.e., prebird), in the absence of a living analogue, a new approach using Strouhal number     was used. Strouhal number is a nondimensional number that describes the relationship between wing-stroke amplitude ( A ), wing-beat frequency ( f ), and flight speed ( U ). Calculations indicated that even moderate wing movements are enough to generate rudimentary thrust and that a propulsive flapping flight-stroke could have evolved via gradual incremental changes in wing movement and wing morphology. More fundamental to the origin of the avian flapping flight-stroke is the question of how a symmetrical forelimb posture—required for gliding and flapping flight—evolved from an alternating forelimb motion, evident in all extant bipeds when running except birds.     

小脑电刺激对家鸽乙酰胆碱酯酶活力的影响

选用成年健康家鸽,对小脑进行连续电刺激后,分别抽取家鸽外周血清及全脑测定乙酰胆碱酯酶活力变化,以探讨鸟类小脑刺激与乙酰胆碱酯酶活力变化之间的相关性.结果 表明:在刺激小脑皮层后,家鸽外周血清中乙酰胆碱酯酶活力显著上升(P＜0.05);而在刺激小脑皮层后,脑组织中乙酰胆碱酯酶活力显著降低(P＜0.05).推测电刺激引起外周组织乙酰胆碱释放,从而引起肌肉强直,血清中胆碱酯酶的活力升高.而电刺激小脑使抑制性神经元功能兴奋,脑中胆碱能神经元功能减弱,乙酰胆碱的释放减少,脑组织中胆碱酯酶的活力降低.     

Spiders fluoresce variably across many taxa

The evolution of fluorescence is largely unexplored, despite the newfound occurrence of this phenomenon in a variety of organisms. We document that spiders fluoresce under ultraviolet illumination, and find that the expression of this trait varies greatly among taxa in this species-rich group. All spiders we examined possess fluorophores in their haemolymph, but bright fluorescence appears to result when a spider sequesters fluorophores in its setae or cuticle. By sampling widely across spider taxa, we determine that fluorescent expression is labile and has evolved multiple times. Moreover, examination of the excitation and emission properties of extracted fluorophores reveals that spiders possess multiple fluorophores and that these differ among some families, indicating that novel fluorophores have evolved during spider diversification. Because many spiders fluoresce in wavelengths visible to their predators and prey (birds and insects), we propose that natural selection imposed by predator-prey interactions may drive the evolution of fluorescence in spiders.     

The evolution of body size in birds. I. Evidence for non-random diversification

The hypothesis that evolution of body size in birds was a random process coupled with an absolute lower boundary on body mass was tested using data on 6217 species of extant birds. The test was based on the fact that subclades within birds that have body masses much larger than this minimum should not have skewed log body mass distributions, while clades close to this boundary should. Bird species were classified into 23 orders suggested by Sibley and Monroe (1988). Thirteen orders that had average log body masses greater than the average for all birds had significantly skewed log body mass distributions. This is inconsistent with the hypothesis that evolution of body size in birds is random, but is constrained only at the smallest body masses. Most orders of birds cannot be considered random samples from the parental distribution of all birds. When the pattern of body mass evolution in birds is reconstructed using an estimate of the phylogenetic relationships among orders, there are many more instances where a large taxon putatively originated from a smaller one than vice versa. The non-random nature of body mass evolution in birds is consistent with models that postulate that evolution is constrained by the ability of individuals to turn resources into offspring.     

The evolution of stereopsis and the Wulst in caprimulgiform birds: a comparative analysis

Owls possess stereopsis (i.e., the ability to perceive depth from retinal disparity cues), but its distribution amongst other birds has remained largely unexplored. Here, we present data on species variation in brain and telencephalon size and features of the Wulst, the neuroanatomical substrate that subserves stereopsis, in a putative sister-group to owls, the order Caprimulgiformes. The caprimulgiforms we examined included nightjars (Caprimulgidae), owlet-nightjars (Aegothelidae), potoos (Nyctibiidae), frogmouths (Podargidae) and the Oilbird (Steatornithidae). The owlet-nightjars and frogmouths shared almost identical relative brain, telencephalic and Wulst volumes as well as overall brain morphology and Wulst morphology with owls. Specifically, the owls, frogmouths and owlet-nightjars possess relatively large brains and telencephalic and Wulst volumes, had a characteristic brain shape and displayed prominent laminae in the Wulst. In contrast, potoos and nightjars both had relatively small brains and telencephala, and Wulst volumes that are typical for similarly sized birds from other orders. The Oilbird had a large brain, telencephalon and Wulst, although these measures were not quite as large as those of the owls. This gradation of owl-like versus nightjar-like brains within caprimulgiforms has significant implications for understanding the evolution of stereopsis and the Wulst both within the order and birds in general.     

Reptilian Cognition: A More Complex Picture via Integration of Neurological Mechanisms,Behavioral Constraints,and Evolutionary Context

Unlike birds and mammals, reptiles are commonly thought to possess only the most rudimentary means of interacting with their environments, reflexively responding to sensory information to the near exclusion of higher cognitive function. However, reptilian brains, though structurally somewhat different from those of mammals and birds, use many of the same cellular and molecular processes to support complex behaviors in homologous brain regions. Here, the neurological mechanisms supporting reptilian cognition are reviewed, focusing specifically on spatial cognition and the hippocampus. These processes are compared to those seen in mammals and birds within an ecologically and evolutionarily relevant context. By viewing reptilian cognition through an integrative framework, a more robust understanding of reptile cognition is gleaned. Doing so yields a broader view of the evolutionarily conserved molecular and cellular mechanisms that underlie cognitive function and a better understanding of the factors that led to the evolution of complex cognition.     

New data on the brain and cognitive abilities of birds

New evidence of functional analogies and homologies of avian and mammalian brains is presented, as is a revised nomenclature of the most important brain structures. Comparative characteristics of the avian brain and criteria for its progressive development in the phylogeny have been considered. We studied the possibility to use Portmann??s index as one of the indicators of brain development in different avian species. We substantiated the necessity to chose for investigation new sets of avian species with medium (Parus caeruleus and Loxia curvirostra) and low (Larus glaucescens) levels of brain complexity to maintain fully valuable grounds for comparing the cognitive abilities in birds. The main experimentally supported proofs of the existence of elementary thinking and some other cognitive functions in the higher birds have been reviewed. The high levels of cognitive processes that underlie the tool using ability in birds, as well as the similarity to those processes in apes, have been demonstrated from the results obtained in the first decade of the 21st century. Comparative studies on proto-instrumental activity confirmed the ability of hooded crows and ravens to find urgent solution of tool-using tasks. Although birds with a medium level of brain complexity display seemingly rational behavior, it is plausible that they use simpler rules being unable to understand the task logic. It was shown that birds of different orders with a high level of brain complexity demonstrate similar dynamics in the development of abstract concepts. Crossbills, which have a medium level of brain complexity, were able to develop the same concepts at a lower level than the corvids; whereas the seagulls and pigeons, which possess a low level of cognitive abilities, were not able to operate any abstractions and were incapable of solving other cognitive tests. The fact that corvids, parrots, and apes have similar abilities to solve some cognitive tasks supports the hypothesis of the convergent evolution of the brain and cognition in birds and primates.     

Is Cooperative Breeding Associated With Bigger Brains? A Comparative Test in the Corvida (Passeriformes)

The cognitive demands of a social existence favour the evolution of relatively large brains and neocortices in primates. Comparable tests of sociality and brain size/structure in birds have not been performed, despite marked similarities in the social systems of birds and mammals. Here, we test whether one aspect of avian sociality, cooperative breeding, is associated with an increase in brain size across 155 species of the passeriform parvorder Corvida. Using conventional and phylogeny‐corrected statistics, we examined the correlated evolution of relative brain size and: the presence/absence of cooperative breeding, percentage of nests that are cooperative and cooperatively breeding group size. Most of the comparisons yielded non‐significant results, which suggests that cooperative breeding is not related to relative brain size in this parvorder. There are a number of potential explanations for our findings. First, changes in brain region size may be correlated with cooperative breeding without affecting overall brain size. Secondly, cooperatively breeding birds might not possess more complex social behaviour than non‐cooperatively breeding birds. Thirdly, relatively large brains might be ancestral in this parvorder. This may predispose them to evolve the range of complex behaviours found in this group, including extreme sociality. Finally, ecological and/or developmental factors might play a more significant role than social behaviour in the diversification of avian brain size. Assessing these alternatives requires more information on the neural and cognitive differences between bird species.     

Evolutionary significance of foregut fermentation in the hoatzin (Opisthocomus hoazin; Aves: Opisthocomidae)

The hoatzin (Opisthocomus hoazin) is the only folivorous bird known to possess extensive fermentation in the crop by mixed bacterial populations. In this work, the digestive tract of the hoatzin was studied morphometrically and microbiologically, and its significance in the evolution of herbivory in vertebrates is discussed. The crop of the hoatzin is already formed in newly hatched chicks, and acquires microbial populations of bacteria and protozoa within the first 2 weeks of life, presumably by inoculation during feeding by adults. Numbers of bacteria and protozoa resemble numbers from the rumen of Ruminantia. The presence of foregut fermentation in this bird demonstrates that this strategy is not an exclusive feature of mammals. Herbivorous mammals developed foregut structures without homologues among other mammals, while the crop of the hoatzin, being homologous to that in other birds, is analogous to the mammalian foregut fermentation chambers. Thus, evolution of foregut fermentation in mammals and birds might be a case of evolutionary convergence.Abbreviations BM body mass - GC gas-liquid-chromatography - SCFA short-chain fatty acid - SE standard error of mean     

Virtual endocasts: an application of computed tomography in the study of brain variation among hyenas

Reliable brain volume measurements are crucial in identifying factors that influence the course of brain evolution. Here, we demonstrate the potential for using virtual endocasts (VEs) to examine inter- and intraspecific variation in brain volume in members of the family Hyaenidae. Total endocranial volume (adjusted for body size) and anterior cerebrum volume (adjusted for endocranial volume) were greater in the spotted hyena, the most gregarious of the species, than in the other hyaenids, all of which are less gregarious. An intraspecific analysis of spotted hyenas revealed that anterior cerebrum volume is significantly larger in males than females, although total endocranial volume does not differ between the sexes. Greater total endocranial and anterior cerebrum volume of spotted hyenas, relative to those of other hyena species, may be related to increased neural processing mediating cognitive demands associated with a complex social life. These data demonstrate that computed tomographic (CT) technology can be used to create VEs in species for which actual brains are rare or unavailable, and suggest that this approach can be applied systematically to explore intra- and interspecies brain variations in studies of brain evolution.     

鸟类线粒体DNA研究概述

线粒体DNA作为理想的分子标记已被广泛用于鸟类种群遗传学和进化遗传学的研究,并取得了许多有意义的结果。本文介绍鸟类线粒体DNA的组成、结构特点及多态性的研究,综述近年来有关鸟类分子进化研究的进展情况,对今后的发展进行了初步的探讨。 Abstract:Mitochondrial DNA as a genetic marker has been successfully applied to the study of molecular evolution of birds.The apparently maternal inheritance of mitochondrial DNA and its fast evolution in primary sequence has made it attractive in population and evolutionary genetics.Mitochondrial DNA of birds displays two characteristics not seen in other vertebrates mtDNA,that is,a novel gene order and the absence of an equivalent to the light-strand replication origin.The research on polymorphism of mtDNA can resolve phylogenies of birds both at lower and higher taxonomic levels.Here we review progress on avian molecular evolution in recent years,and make preliminary studies of the development in this field.     

Tasting the difference: do multiple defence chemicals interact in Müllerian mimicry?

Müllerian mimicry, where two unpalatable species share a warning pattern, is classically believed to be a form of mutualism, where the species involved share the cost of predator education. The evolutionary dynamics of Müllerian mimicry have recently become a controversial subject, after mathematical models have shown that if minor alterations are made to assumptions about the way in which predators learn and forget about unpalatable prey, this textbook case of mutualism may not be mutualistic at all. An underlying assumption of these models is that Müllerian mimics possess the same defence chemical. However, some Müllerian mimics are known to possess different defence chemicals. Using domestic chicks as predators and coloured crumbs flavoured with either the same or different unpalatable chemicals as prey, we provide evidence that two defence chemicals can interact to enhance predator learning and memory. This indicates that Müllerian mimics that possess different defence chemicals are better protected than those that share a single defence chemical. These data provide insight into how multiple defence chemicals are perceived by birds,and how they influence the way birds learn and remember warningly coloured prey. They highlight the importance of considering how different toxins in mimicry rings can interact in the evolution and maintenance of Müllerian mimicry and could help to explain the remarkable variation in chemical defences found within and between species.     

Perforin evolved from a gene duplication of MPEG1, followed by a complex pattern of gene gain and loss within Euteleostomi

ABSTRACT: BACKGROUND: The pore-forming protein perforin is central to the granule-exocytosis pathway used by cytotoxic lymphocytes to kill abnormal cells. Although this mechanism of killing is conserved in bony vertebrates, cytotoxic cells are present in other chordates and invertebrates, and their cytotoxic mechanism has not been elucidated. In order to understand the evolution of this pathway, here we characterize the origins and evolution of perforin. RESULTS: We identified orthologs and homologs of human perforin in all but one species analysed from Euteleostomi, and present evidence for an earlier ortholog in Gnathostomata but not in more primitive chordates. In placental mammals perforin is a single copy gene, but there are multiple perforin genes in all lineages predating marsupials, except birds. Our comparisons of these many-to-one homologs of human perforin show that they mainly arose from lineagespecific gene duplications in multiple taxa, suggesting acquisition of new roles or different modes of regulation. We also present evidence that perforin arose from duplication of the ancient MPEG1 gene, and that it shares a common ancestor with the functionally related complement proteins. CONCLUSIONS: The evolution of perforin in vertebrates involved a complex pattern of gene, as well as intron, gain and loss. The primordial perforin gene arose at least 500 million years ago, at around the time that the major histocompatibility complex-T cell receptor antigen recognition system was established. As it is absent from primitive chordates and invertebrates, cytotoxic cells from these lineages must possess a different effector molecule or cytotoxic mechanism.     

Succinic Dehydrogenase Distribution in the Pectoralis Muscle of Several East African Birds

The distribution of succinic dehydrogenase activity was investigated in the pectoralis muscle of thirteen East African birds, representing five Orders. It was found that the pectoralis muscle of the most primitive birds studied (Galliformes) contained all “white” muscle fibres whereas the more advanced birds (Passeriformes) had all “red” muscle fibres. Intermediate Orders had mostly a mixture of red and white muscle fibres. There also appeared to be a direct relationship between body size and average muscle fibre size. However, it was concluded that the most important factor in relation to the muscle structure is the bird's mode of flight. The relationship with the degree of evolution and body size only held true in so far as the birds which had developed the facility for sustained flight, by increasing their red muscle fibre content, were also smaller in size and constituted the more “evolved” Orders of birds. In support of this it was noted that migratory birds (i.e. engaging in sustained flight) from more primitive Orders also had a high red muscle fibre content in their pectoralis muscles.     

Multiple Ornaments in Male Northern Cardinals, Cardinalis cardinalis, as Indicators of Condition

Investigations of male ornaments in the context of sexual selection have tended to focus on single ornaments, although many species of birds possess multiple ornaments. Understanding the evolution of multiple ornaments requires knowledge of correlations among ornaments in the same individual and the extent to which ornament expression reflects individual condition and behavior. Variation in four male ornaments in socially monogamous, biparental northern cardinals (Cardinalis cardinalis) was related to body size, indices of condition, level of paternal care, and reproductive success. Redness of breast plumage positively predicted body size and negatively predicted nestling feeding rate. Bill color predicted current body condition, with birds with redder bills in better condition. Birds with smaller black face masks had greater reproductive success. These results are consistent with the hypothesis that different ornaments in male cardinals provide information on different aspects of condition and behavior.     

Avian reticuloendotheliosis viruses: evolutionary linkage with mammalian type C retroviruses.

Reticuloendotheliosis viruses have been shown to be causative of tumors in a variety of avian species. The major structural protein of these non-genetically transmitted viruses is demonstrated to possess antigenic determinants common to those of all known mammalian type C viruses. These findings establish a mammalian origin for this oncogenic avian retrovirus group. None of the known mammalian type C virus groups demonstrated a closer immunological relationship to avian reticuloendotheliosis viruses. These results suggest that reticuloendotheliosis viruses have been non-genetically transmitted for a long period of evolution or that these viruses may have arisen by relatively recent infection of birds with an as yet undiscovered mammalian type C retrovirus.     

Carotenoids modulate the trade-off between egg production and resistance to oxidative stress in zebra finches

The allocation of resources to reproduction and survival is a central question of studies of life history evolution. Usually, increased allocation to current reproduction is paid in terms of reduced future reproduction and/or decreased survival. However, the proximal mechanisms underlying the cost of reproduction are poorly understood. Recently, it has been shown that increased susceptibility to oxidative stress might be one of such proximate links between reproduction and self-maintenance. Organisms possess a range of antioxidant defenses, including endogenously produced molecules (e.g., enzymes) and compounds ingested with food (e.g., carotenoids). If reproductive effort increases the production of reactive oxygen species, the availability of antioxidant defenses may partly or fully counteract the free-radical damages. One could, therefore, expect that the trade-off between reproduction and oxidative stress is modulated by the availability of antioxidant defenses. We tested this hypothesis in zebra finches. We manipulated reproductive effort by either allowing or preventing pairs to breed. Within each breeding or non-breeding group, the availability of antioxidant compounds was manipulated by supplementing or not supplementing the drinking water with carotenoids. We found that although birds in the breeding and non-breeding groups did not differ in their resistance to oxidative stress (the breakdown of red blood cells submitted to a controlled free-radical attack), one aspect of breeding effort (i.e., the number of eggs laid by birds in both breeding and non-breeding groups) was negatively correlated with resistance to oxidative stress only in birds that did not benefit from a carotenoid-supplemented diet. This result therefore suggests that carotenoid availability can modulate the trade-off between reproduction and resistance to oxidative stress.     

Adaptive molecular evolution of HINTW,a female-specific gene in birds

It is well established that many genes on the male-specific Y chromosome of organisms such as mammals are involved in male reproduction and may evolve rapidly because of positive selection on male reproductive traits. In contrast, very little is known about the function and evolution of W-linked genes restricted to the female genome of organisms with female heterogamety. For birds (males ZZ, females ZW), only one W-linked gene (HINTW) is sufficiently different from its Z-linked homolog to indicate a female-specific function. Here, we report that HINTW shows evidence of adaptive molecular evolution, implying strong positive selection for new functional properties in female birds. Moreover, because HINTW is expressed in the gonads of female birds just before sexual differentiation and is thus a candidate for sex determination, it suggests adaptive evolution related to female development. This provides the first example of Darwinian evolution of a gene restricted to the female genome of any organism. Given that HINTW exists in multiple copies on W, similar to some testis-specific genes amplified on mammalian Y, avian HINTW may thus potentially represent a female parallel to the organization and evolution of Y chromosome genes involved in male reproduction and development.