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Exaptation and Evolutionary Adaptation in Nociceptor Mechanisms Driving Persistent Pain. Brain Behav. Evol. (IF 1.7) Pub Date : 2023-11-30 Edgar T Walters
BACKGROUND Several evolutionary explanations have been proposed for why chronic pain is a major clinical problem. One is that some mechanisms important for driving chronic pain, while maladaptive for modern humans, were adaptive because they enhanced survival. Evidence is reviewed for persistent nociceptor hyperactivity (PNH), known to promote chronic pain in rodents and humans, being an evolutionarily
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Hippocampal, Whole Midbrain, Red Nucleus, and Ventral Tegmental Area Volumes Are Increased by Selective Breeding for High Voluntary Wheel-Running Behavior. Brain Behav. Evol. (IF 1.7) Pub Date : 2023-08-21 Margaret P Schmill,Zoe Thompson,Daisy Lee,Laurence Haddadin,Shaarang Mitra,Raymond Ezzat,Samantha Shelton,Phillip Levin,Sogol Behnam,Kelly J Huffman,Theodore Garland
Uncovering relationships between neuroanatomy, behavior, and evolution are important for understanding the factors that control brain function. Voluntary exercise is one key behavior that both affects, and may be affected by, neuroanatomical variation. Moreover, recent studies suggest an important role for physical activity in brain evolution. We used a unique and ongoing artificial selection model
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Sex Differences in the Neural Song Circuit and Its Relationship to Song Acoustic Complexity in House Wrens (Troglodytes aedon). Brain Behav. Evol. (IF 1.7) Pub Date : 2023-07-24 Cara A Krieg,Juli Wade
The song circuit in passerine birds is an outstanding model system for understanding the relationship between brain morphology and behavior, in part due to varying degrees of sex differences in structure and function across species. House wrens (Troglodytes aedon) offer a unique opportunity to advance our understanding of this relationship. Intermediate sex differences in song rate and complexity exist
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The relationship between cognition and brain size or neuron number. Brain Behav. Evol. (IF 1.7) Pub Date : 2023-07-24 Andrew B Barron,Faelan Mourmourakis
The comparative approach is a powerful way to explore the relationship between brain structure and cognitive function. Thus far the field has been dominated by the assumption that a bigger brain somehow means better cognition. Correlations between differences in brain size or neuron number between species and differences in specific cognitive abilities exist, but these correlations are very noisy.
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Conservation and Diversification of Pallial Cell Types across Vertebrates: An Evo-Devo Perspective. Brain Behav. Evol. (IF 1.7) Pub Date : 2023-06-28 Shreyas M Suryanarayana,Dhananjay Huilgol
As the highest center of sensory processing, initiation, and modulation of behavior, the pallium has seen prominent changes during the course of vertebrate evolution, culminating in the emergence of the mammalian isocortex. The processes underlying this remarkable evolution have been a matter of debate for several centuries. Recent studies using modern techniques in a host of vertebrate species are
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Mapping human brain pathways: challenges and opportunities in the integration of scales Brain Behav. Evol. (IF 1.7) Pub Date : 2023-03-27
The human brain is composed of a complex web of pathways. Diffusion magnetic resonance (MR) tractography relies on the principle of diffusion to reconstruct brain pathways. Its tractography is broadly applicable to a range of problems as it is amenable to being studied in individuals of any age and from any species. However, it is well-known that this technique can generate biologically implausible
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Evolution of Local Circuit Neurons in Two Sensory Thalamic Nuclei in Amniotes Brain Behav. Evol. (IF 1.7) Pub Date : 2023-03-27 Michael B. Pritz
Local circuit neurons are present in the thalamus of all vertebrates where they are considered inhibitory. They play an important role in computation and influence the transmission of information from the thalamus to the telencephalon. In mammals, the percentage of local circuit neurons in the dorsal lateral geniculate nucleus remains relatively constant across a variety of species. In contrast, the
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The Diversity of the Brains of Ray-Finned Fishes Brain Behav. Evol. (IF 1.7) Pub Date : 2023-03-22
Brains are very plastic both in response to phenotypic diversity and to larger evolutionary trends. Differences between taxa cannot be easily attributed to either factors. Comparative morphological data on higher taxonomic levels is scarce especially in ray-finned fishes. Here we show the great diversity of brain areas of more than 150 species of ray-finned fishes by volumetric measurements using block
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Studying the brain monoaminergic systems and neurotrophic factors in minipigs with high and low tolerance to the presence of human Brain Behav. Evol. (IF 1.7) Pub Date : 2023-03-13 Darya Bazovkina, Nina B. Illarionova, Elizabeth A. Kulikova, Ekaterina Yu. Bazhenova, Nadezhda A. Sinyakova, Nikita V. Khotskin, Sergey Nikitin, Vasily S. Lankin, Elena E. Terenina, Oleg V. Trapezov, Aleksandr V. Kulikov
Here, we present the first evidence for brain adaptation in pigs tolerant to the human presence, as a behavioral trait favoring domestication. The study was carried out on minipiglets from population bred in the Institute of Cytology and Genetics (Novosibirsk, Russia). We compared the behavior, metabolism of monoaminergic neurotransmitter systems and functional activity of the hypothalamic-pituitary-adrenal
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Sex and season affect cortical volumes in free-living Western fence lizards, Sceloporus occidentalis Brain Behav. Evol. (IF 1.7) Pub Date : 2023-02-16 Morgan B Jude, Christine. R. Strand
The hippocampus plays an important role in spatial navigation and spatial learning across a variety of vertebrate species. Sex and seasonal differences in space use and behavior are known to affect hippocampal volume. Similarly, territoriality and differences in home range size are known to affect the volume of the reptile hippocampal homologues, the medial and dorsal cortices (MC, DC). However, studies
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Retinal ganglion cell topography and spatial resolution in three Indian pteropodid bats Brain Behav. Evol. (IF 1.7) Pub Date : 2022-12-29 Baheerathan Murugavel, Mindaugas Mitkus, Hema Somanathan, Almut Kelber
Pteropodidae is the only phytophagous bat family that predominantly depends on visual and olfactory cues for orientation and foraging. During daytime, pteropodids of different species roost in sites with varying light exposure. Pteropodids have larger eyes relative to body size than insectivorous bats. Retinal topography has been studied in less than 10% of the approximately 200 pteropodid species
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Endocranial casts of Camelops hesternus and Palaeolama sp., new insights into the recent history of the camelid brain Brain Behav. Evol. (IF 1.7) Pub Date : 2022-12-27 Ana M. Balcarcel, Dylan Bastiaans, Maeva J Orliac
Endocranial casts are capable of capturing the general brain form in extinct mammals due to the high fidelity of the endocranial cavity and the brain in this clade. Camelids, the clade including extant camels, llamas, and alpacas, today display high levels of gyrification and brain complexity. The evolutionary history of the camelid brain has been described as involving unique neocortical growth dynamics
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Climate Change Influences Brain Size in Humans Brain Behav. Evol. (IF 1.7) Pub Date : 2022-12-27
Brain size evolution in hominins constitutes a crucial evolutionary trend, yet the underlying mechanisms behind those changes are not well understood. Here, climate change is considered as an environmental factor using four paleoclimate records testing temperature, humidity, and precipitation against changes to brain size in 298 Homo specimens over the past fifty thousand years. Across regional and
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Molecular diversity of neuron types in the salamander amygdala and implications for amygdalar evolution Brain Behav. Evol. (IF 1.7) Pub Date : 2022-12-27 Astrid Deryckere, Jamie Woych, Eliza C B Jaeger, Maria Antonietta Tosches
The amygdala is a complex brain structure in the vertebrate telencephalon, essential for regulating social behaviors, emotions and (social) cognition. In contrast to the vast majority of neuron types described in the many nuclei of the mammalian amygdala, little is known about the neuronal diversity in non-mammals, making reconstruction of its evolution particularly difficult. Here, we characterize
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Cytoarchitecture of Mudskipper (Boleophthalmus pectinirostris) Brain Brain Behav. Evol. (IF 1.7) Pub Date : 2022-12-21 Ke Jiang, Ke Wei, Shi Xi Chen, Jing Huang
Mudskippers are intertidal burrowing fish with unique living habits. So far, studies on the cytoarchitecture of brain in fish with such behaviors remain limited. Therefore, documenting the neuroanatomy of this animal is of interest because of its unique characteristics. In this study, we examined the cytoarchitecture of the mudskipper (Boleophthalmus pectinirostris) brain and investigated whether it
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Cerebellar inputs in the American alligator (Alligator mississippiensis) Brain Behav. Evol. (IF 1.7) Pub Date : 2022-12-01 Cristián Gutiérrez-Ibáñez , Lutz Kettler, Madison C Pilon, Catherine E. Carr, Douglas R. Wylie
Crocodilians (alligators, crocodiles and gharials) are the closet living relatives to birds, and as such, represent a key clade to understand the evolution of the avian brain. However, many aspects of crocodilians neurobiology remain unknown. In this paper we address an important knowledge gap as there are no published studies of cerebellar connections in any crocodilian species. We used injections
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Acknlowledgement to Reviewers Brain Behav. Evol. (IF 1.7) Pub Date : 2022-11-23
Brain Behav Evol 2022;97:369
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Evolution and development of amygdala subdivisions: pallial, subpallial and beyond Brain Behav. Evol. (IF 1.7) Pub Date : 2022-10-20 Loreta Medina, Antonio Abellán, Lorena Morales, Alessandra Pross, Alek H Metwalli, Alba González-Alonso, Júlia Freixes, Ester Desfilis
The amygdala is a central node in functional networks regulating emotions, social behavior and social cognition. It develops in the telencephalon and includes pallial and subpallial parts, but these are extremely complex with multiple subdivisions, cell types and connections. The homology of the amygdala in non-mammals is highly controversial, especially for the pallial part, and we are still far from
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Evolution and development of amygdala subdivisions: pallial, subpallial and beyond Brain Behav. Evol. (IF 1.7) Pub Date : 2022-10-20
The amygdala is a central node in functional networks regulating emotions, social behavior and social cognition. It develops in the telencephalon and includes pallial and subpallial parts, but these are extremely complex with multiple subdivisions, cell types and connections. The homology of the amygdala in non-mammals is highly controversial, especially for the pallial part, and we are still far from
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The 42nd Annual Meeting of the J.B. Johnston Club for Evolutionary Neuroscience and the 34th Annual Karger Workshop in Evolutionary Neuroscience Brain Behav. Evol. (IF 1.7) Pub Date : 2022-08-23 Muhammad Spocter
The 2022 meetings of the J.B. Johnston Club for Evolutionary Neuroscience and Karger Workshop in Evolutionary Neuroscience will be held immediately before the annual meeting of the Society for Neuroscience on Thursday, November 10 (the Karger Workshop), and Friday, November 11 (the regular JBJC meeting). Both meetings will take place at the Horton Grand Hotel, San Diego CA, USA. This year’s Karger
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The 42nd Annual Meeting of the J.B. Johnston Club for Evolutionary Neuroscience and the 34th Annual Karger Workshop in Evolutionary Neuroscience Brain Behav. Evol. (IF 1.7) Pub Date : 2022-08-23
The 2022 meetings of the J.B. Johnston Club for Evolutionary Neuroscience and Karger Workshop in Evolutionary Neuroscience will be held immediately before the annual meeting of the Society for Neuroscience on Thursday, November 10 (the Karger Workshop), and Friday, November 11 (the regular JBJC meeting). Both meetings will take place at the Horton Grand Hotel, San Diego CA, USA. This year’s Karger
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Mapping the pattern of essential neuroendocrine cells related to puberty and VA opsin expression provides further insight in the photoreceptive regulation of the BPG axis in Atlantic salmon (Salmo salar) Brain Behav. Evol. (IF 1.7) Pub Date : 2022-08-03 Christine Horne, Jon Vidar Helvik, Mitchell Stewart Fleming, Per Gunnar Fjelldal, Mariann Eilertsen
In Atlantic salmon (Salmo salar) seasonal photoperiod is shown to regulate the onset of sexual maturation, yet which brain region(s) are involved and how light information impacts the neuroendocrine system are still not fully understood in teleosts. Detailed knowledge about the photoperiodic regulation of maturation in fish is still missing. In birds, it is shown that gonadotrophin releasing hormone
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Mapping the Pattern of Essential Neuroendocrine Cells Related to Puberty and VA Opsin Expression Provides Further Insight in the Photoreceptive Regulation of the Brain-Pituitary-Gonadal Axis in Atlantic Salmon (Salmo salar) Brain Behav. Evol. (IF 1.7) Pub Date : 2022-08-03
In Atlantic salmon (Salmo salar), seasonal photoperiod is shown to regulate the onset of sexual maturation, yet which brain region(s) is involved, and how light information impacts the neuroendocrine system are still not fully understood in teleosts. Detailed knowledge about the photoperiodic regulation of maturation in fish is still missing. In birds, it is shown that gonadotropin-releasing hormone
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Author Index/Subject Index Vol. 97, No. 3-4, 2022 Brain Behav. Evol. (IF 1.7) Pub Date : 2022-07-01
Brain Behav Evol 2022;97:253
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Author Index/Subject Index Vol. 97, No. 3-4, 2022 Brain Behav. Evol. (IF 1.7) Pub Date : 2022-07-01
Brain Behav Evol 2022;97:253
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The Everted Amygdala of Ray-Finned Fish—Zebrafish Makes a Case Brain Behav. Evol. (IF 1.7) Pub Date : 2022-06-27 Thomas Mueller
The amygdala, a complex array of nuclei in the forebrain, controls emotions and emotion-related behaviors in vertebrates. Current research aims to understand the amygdala’s evolution in ray-finned fish such as zebrafish because of the region’s relevance for social behavior and human psychiatric disorders. Clear-cut molecular definitions of the amygdala and its evolutionary-developmental relationship
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The Everted Amygdala of Ray-Finned Fish: Zebrafish Makes a Case Brain Behav. Evol. (IF 1.7) Pub Date : 2022-06-27
The amygdala, a complex array of nuclei in the forebrain, controls emotions and emotion-related behaviors in vertebrates. Current research aims to understand the amygdala’s evolution in ray-finned fish such as zebrafish because of the region’s relevance for social behavior and human psychiatric disorders. Clear-cut molecular definitions of the amygdala and its evolutionary-developmental relationship
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The neuromeric/prosomeric model in teleost fish neurobiology Brain Behav. Evol. (IF 1.7) Pub Date : 2022-06-21 Mario F. Wullimann
The neuromeric/prosomeric model has been rejuvenated by Puelles and Rubenstein (1993). Here, its application to the (teleostean) fish brain is detailed beginning with a historical account. The second part addresses three main issues with particular interest for fish neuroanatomy and looks at the impact of the neuromeric model on their understanding. The first one is the occurence of four early migrating
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The Neuromeric/Prosomeric Model in Teleost Fish Neurobiology Brain Behav. Evol. (IF 1.7) Pub Date : 2022-06-21
The neuromeric/prosomeric model has been rejuvenated by Puelles and Rubenstein [Trends Neurosci. 1993;16(11):472–9]. Here, its application to the (teleostean) fish brain is detailed, beginning with a historical account. The second part addresses three main issues with particular interest for fish neuroanatomy and looks at the impact of the neuromeric model on their understanding. The first one is the
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Author Index/Subject Index Brain Behav. Evol. (IF 1.7) Pub Date : 2022-06-09
Brain Behav Evol 2022;97:121
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Acknowledgement to Reviewers Brain Behav. Evol. (IF 1.7) Pub Date : 2022-06-09
Brain Behav Evol 2022;97:122
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Author Index/Subject Index Brain Behav. Evol. (IF 1.7) Pub Date : 2022-06-09
Brain Behav Evol 2022;97:121
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Acknowledgement to Reviewers Brain Behav. Evol. (IF 1.7) Pub Date : 2022-06-09
Brain Behav Evol 2022;97:122
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Author Index/Subject Index Brain Behav. Evol. (IF 1.7) Pub Date : 2022-05-31
Brain Behav Evol 2021;96:373
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Comparative neuroanatomy of the mechanosensory subgenual organ complex in the Peruvian stick insect, Oreophoetes peruana Brain Behav. Evol. (IF 1.7) Pub Date : 2022-06-02 Johannes Strauß
The subgenual organ complex in the leg of Polyneoptera (Insecta) consist of several chordotonal organs specialised to detect mechanical stimuli from substrate vibrations and airborne sound. In stick insects (Phasmatodea), the subgenual organ complex contains the subgenual organ and the distal organ located distally to the subgenual organ. The subgenual organ is a highly sensitive detector for substrate
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Publications of Luis Puelles in Developmental and Comparative Neurobiology Brain Behav. Evol. (IF 1.7) Pub Date : 2022-05-31
Brain Behav Evol 2021;96:355–363
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Selected Publications of Agustín González in Comparative Neuroscience Brain Behav. Evol. (IF 1.7) Pub Date : 2022-05-31
Brain Behav Evol 2021;96:364–371
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Contents Vol. 96, 2021 Brain Behav. Evol. (IF 1.7) Pub Date : 2022-05-31
Brain Behav Evol 2021;96:I–IV
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Organizational conservation and flexibility in the evolution of birdsong and avian motor control Brain Behav. Evol. (IF 1.7) Pub Date : 2022-05-27 Bradley M Colquitt
Birds and mammals have independently evolved complex behavioral and cognitive capabilities yet have markedly different brain structures. An open question is to what extent, despite these differences in anatomy, birds and mammals have evolved similar neural solutions to complex motor control and at what level of organization these similarities might lie. Courtship song in songbirds, a learned motor
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ANALYSIS OF THE PALLIAL AMYGDALA IN ANURANS: DERIVATIVES AND CELLULAR COMPONENTS Brain Behav. Evol. (IF 1.7) Pub Date : 2022-05-25 Sara Jiménez, Nerea Moreno
The amygdaloid complex plays a crucial role in socio-emotional conduct, learning, survival, and reproductive behaviors. It is constituted by a set of nuclei presenting a great cellular heterogeneity and embryonic origin diversity (pallial, subpallial and even extra-telencephalic). In the last two decades, the tetrapartite pallial paradigm defined the pallial portion of the amygdala as a derivative
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Higher rate of male sexual displays correlates with larger ventral posterior amygdala volume and neuron soma volume in wild-caught common side-blotched lizards, Uta stansburiana Brain Behav. Evol. (IF 1.7) Pub Date : 2022-05-10
Several areas of the vertebrate brain are involved in facilitating and inhibiting the production of sexual behaviors and displays. In the laboratory, a higher rate of sexual displays is correlated with a larger ventral posterior amygdala (VPA), an area of the brain involved in the expression of sexual display behaviors, as well as larger VPA neuronal somas. However, it remains unclear if individuals
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Acknowledgement to Guest Editors, Authors, and Reviewers Brain Behav. Evol. (IF 1.7) Pub Date : 2022-04-12 Philippe Vernier, Kara Yopak, Georg F. Striedter, Walter Wilczynski, Blinda McClelland, Ann B. Butler, Eric J. Vallender, Andrew N. Iwaniuk, Roger L. Reep, Sunny K. Boyd, Eliot Brenowitz, Alice Powers, N.J. Strausfeld, Günther K.H. Zupanc, Jon H. Kaas, Naoyuki Yamamoto, T.M. Preuss, Anton Reiner, Robert Huber, Mario F. Wullimann, Karen A. Mesce, Susan E. Fahrbach, Catherine A. Marler, Horst Bleckmann
Brain Behav Evol
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Epilogue: A Tribute to Luis Puelles and Agustín González. Brain Behav. Evol. (IF 1.7) Pub Date : 2022-04-06 Philippe Vernier
The scientific context of the special issue dedicated to Luis Puelles and Agustin Gonzalez is given in this short epilogue.
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In Memory of Walter Wilczynski: Neuroethologist, Mentor, Role Model and Friend. Brain Behav. Evol. (IF 1.7) Pub Date : 2022-04-06 Blinda E McClelland,Kathleen S Lynch
Introduction to Walter Wilczynski Special BBE Edition.
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Evolution of developmental timing as a driving force of brain diversity Brain Behav. Evol. (IF 1.7) Pub Date : 2022-03-28 Rodrigo Suárez, Andrew C Halley
The question of how complex traits originate and diversify has marveled naturalists for millennia. From the notion of development as a series of transformations beyond ‘pre-formed’ growth by Aristotle, to von Baer’s recognition of phylogenetic differentiation that set the foundations of modern evo-devo thinking, a central theme has been the nature of biological change (and conservation) across temporal
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Neurogenetic heterochrony in chick, lizard and rat mapped with wholemount AChE and the prosomeric model. Brain Behav. Evol. (IF 1.7) Pub Date : 2022-03-23 José A Amat, Margaret Martínez-de-la-Torre, Carmen María Trujillo, Bárbara Fernández, Luis Puelles
In the developing brain, the phenomenon of neurogenesis is manifested heterotopically, that is, much the same neurogenetic steps occur at different places with a different timetable. This is due apparently to early molecular regionalization of the neural tube wall in the anteroposterior and dorsoventral dimensions, in a checkerboard pattern of more or less deformed quadrangular histogenetic areas.
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Signaler’s vasotocin alters relationship between responder’s forebrain catecholamines and communication behavior in lizards (Anolis carolinensis) Brain Behav. Evol. (IF 1.7) Pub Date : 2022-03-23 Stephanie M Campos, Anastasia Erley, Zoha Ashraf, Walter Wilczynski
Dynamic fluctuations in the distribution of catecholamines across the brain modulate the responsiveness of vertebrates to social stimuli. Previous work demonstrates that green anoles (Anolis carolinensis) increase chemosensory behavior in response to males treated with exogenous arginine vasotocin (AVT), but the neurochemical mechanisms underlying this behavioral shift remains unclear. Since central
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Relative brain volume of carnivorans has evolved in correlation with environmental and dietary variables differentially among clades Brain Behav. Evol. (IF 1.7) Pub Date : 2022-03-02 Leigha M. Lynch, Kari L. Allen
Carnivorans possess relatively large brains compared to most other mammalian clades. Factors like environmental complexity (Cognitive Buffer Hypothesis) and diet quality (Expensive-Tissue Hypothesis) have been proposed as mechanisms for encephalization in other large-brained clades. We examine whether the Cognitive Buffer and Expensive-Tissue Hypotheses account for brain size variation within Carnivora
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Luis Puelles, the learned neuroembryologist and comparative neurobiologist Brain Behav. Evol. (IF 1.7) Pub Date : 2022-02-28 Salvador Martinez
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Role of melatonin in temperature-induced activation of the neuroendocrine reproductive axis in garter snakes. Brain Behav. Evol. (IF 1.7) Pub Date : 2022-02-25 Treven J. Winters, Stephanie Martin, Holden Anderson, Nichole D. Procter, Deborah I. Lutterschmidt
An animal’s ability to respond optimally to changing environmental conditions is paramount to successfully reproducing and thus maximizing fitness. Studies on photoperiod-induced changes in neural thyroid hormone metabolism have conclusively linked environmental cues to the neuroendocrine reproductive axis of birds and mammals. Whether this conserved mechanism also transduces changes in environmental
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The tempo of mammalian embryogenesis: variation in the pace of brain and body development Brain Behav. Evol. (IF 1.7) Pub Date : 2022-02-21 Andrew C Halley
Why do some species develop rapidly, while others develop slowly? Mammals are highly variable in the pace of growth and development over every stage of ontogeny, and this basic variable – the pace of ontogeny – is strongly associated with a wide range of phenotypes in adults, including allometric patterns of brain and body size as well as the pace of neurodevelopment. This analysis describes variation
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Color discrimination provides insight into the relationship between personality cognition and brain morphology in the of western mosquitofish’s (Gambusia affinis) Brain Behav. Evol. (IF 1.7) Pub Date : 2022-02-21 Mengyu Liu, Yanqiu Liu, He Wang, Jia Jia, Kai Liu
Recent studies on the differences in cognitive ability between individuals focused on two aspects: one is whether the individual differences in cognitive ability are related to brain size, the other is whether they pertain to certain personality traits. To explore these two hypotheses, we tested the personality traits, cognitive abilities and brain volumes of western mosquitofish (Gambusia affinis)
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Peripheral auditory system divergence does not explain species differences in call preference Brain Behav. Evol. (IF 1.7) Pub Date : 2022-02-11 Kim L. Hoke, Jakob Christensen-Dalsgaard, Molly C. Womack
Receiver sensory systems have long been cited as an important source of variation in mate preferences that could lead to signal diversification and behavioral isolation between lineages, with a general concept that animals prefer the most conspicuous signals. The matched filter hypothesis posits that tuning of the frog peripheral auditory system matches dominant frequencies in advertisement calls used
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Phenotypic alterations in cortical organization and connectivity across different time scales Brain Behav. Evol. (IF 1.7) Pub Date : 2022-02-03 Mackenzie Englund, Leah Krubitzer
In the following review, we describe the types of phenotypic changes to the neocortex that occur over the longer time scale of evolution, and over the shorter time scale of an individual lifetime. To understand how phenotypic variability emerges in the neocortex, it is important to consider the cortex as part of an integrated system of the brain, the body, the environment in which the brain and body