This paper is a tribute to outstanding evolutionary biologist Dmitri Konstantinovich Belyaev in connection with the forthcoming centenary of his birth. His work on variation-related mechanisms in animal domestication made this process much faster. Belyaev is at length described as a person, as a scientist and as the organizer of the Institute of Cytology and Genetics of the Siberian Branch of the USSR Academy of Sciences. The author had for many years worked as Belyaev’s Deputy for Science, and, when Dmitri Konstantinovich passed away, had for 22 years headed the Institute of Cytology and Genetics. Belyaev’s life was not easy: he was a priest’s son and an “enemy of the people”’s brother. He was in the battlefield all through the Great Patriotic War and became a man of steel. His struggle for the restoration of genetics in the country, his commitment to setting up and maintaining relationships between masters and followers and many more other aspects of his life are considered. The global importance of the “master – follower” paradigm as a basis of the continuity of generations is underlined and re-underlined, for this is a very special point in the scientific community, especially when new schools of science emerge.

The results of a 35-year selection of foxes for aggressive response to humans are reported. Averaged estimates of the phenotypic manifestation of aggressiveness in all selection generations are presented. The dynamics of these estimates shows that the phenotypic response to the selection was obvious only in the first 12 generations. Subsequent selection did not alter the mean aggressiveness score. Analysis of variance was performed for the intergroup variability (among descendants of different mothers) and intragroup variability (among the offspring within a family). The intragroup variability was constantly low. Most likely, the trait is stabilized by maternal prenatal and early neonatal factors. The general tendency in the dynamics of intergroup variability is that it does not decrease over time during selection, no matter how long the population has been under it. It follows from the statistical indices of the phenotypic similarity between parents and offspring that additive interactions are insufficient for the explanation of the persisting variability. The contribution of epistatic interactions is not ruled out, though. Emphasis is laid on the correlated consequences of the selection for aggressiveness and their coordination with the consequences of the selection in the opposite direction, for elimination of aggressive response to humans, or for tameness. The parallelism of correlated changes in the selection in contrasting directions is illustrated by the examples of some physiological and morphological traits. The phenomenon is discussed in the light of classical notions of the resource of cryptic genetic variation and the role of selection in its phenotypic manifestation. Its interpretation also invokes molecular data pointing that some genetic pathways may regulate parameters of both aggression and tameness and that the selection processes in both directions may have some genetic targets in common.

This paper systematizes and generalizes a research cycle devoted to studying the acoustics and vocal behaviour of silver foxes that differ in their tolerance to humans. The research revealed that 50-year selection for tameness toward people resulted in selective use by Tame foxes toward humans of two call types, pant and cackle. At the same time, the selected for aggression toward people Aggressive foxes and the non-selected for behaviour Control foxes, selectively use toward humans cough and snort. Thus, call types representing vocal indicators of friendly and aggressive behaviour of foxes toward humans have been revealed by the research. Nevertheless, experimental domestication did not change vocal behaviour of foxes toward conspecifics; all three strains did not differ by their vocal behaviour toward same-strain silver foxes. Relationship has been investigated between vocal behaviour and degree of tolerance toward people for hybrids between Tame and Aggressive foxes and for backcrosses to Tame and Aggressive foxes. Effect was estimated between fox sex and the degree of human impact on focal fox for variables of fox vocal behaviour. The research revealed the universal for mammals vocal indicators of emotional arousal that are independent of the emotional valence. Characteristics of vocal behaviour that are related with positive and negative emotional valence have been revealed. A simple and effective method for estimating animal discomfort based on ”joint calls” that takes into account the characteristics of all calls irrespective of their acoustic structure has been revealed. The obtained results provide a basis for further comparative studies of the acoustic structure and vocal behaviour for other taxa of the genus Vulpes and the related canid genera (Canis, Cuon, Lycaon).

The domestication of animals was crucial for the civilization evolutionary experiment. The main characteristic of domesticated animal is tame behavior towards man. D.K. Belyaev suggested that the domestication was carried out by man of the Stone Age as unintended selection of nonaggressive animals. This hypothesis was confirmed at the Institute of Cytology and Genetics, Novosibirsk, by unique long-lasting selection of silver foxes, Norway rats and American minks for nonaggressive behavior towards man. The review concentrates on the role of the brain neurotransmitters in the mechanism of animal domestication. Specifically, it describes the effect of selective breeding for nonaggressive behavior on the brain neurotransmitter serotonin (5-HT) system. Our data showed that produced by domestication the conversion of wild, aggressive animal to non-aggressive towards man is associated with changes in the expression of key molecular members determining the brain 5-HT system functioning. Significant difference between aggressive and nonaggressive animals in the 5-HT metabolism, in the expression of principle metabolic enzymes and 5-HT receptors was shown in different species of mammals (silver foxes, Norway rats and American minks). Higher concentration of 5-HT and its metabolite, 5-hydroxyindoleacetic acid, increased activity of the principle enzyme in 5-HT synthesis (tryptophan hydroxylase-2) and decreased activity of the principle enzyme in 5-HT degradation (monoamine oxidase A) was shown in the midbrain of domesticated silver foxes and rats. The expression of the 5-HTA receptor gene, the density and functional activity of 5-HTA receptors in nonaggressive rats were increased. The review provides converging lines of evidence for our hypothesis that the changes in the brain 5-HT contributes to a mechanism underlying correlative patterns of domestication, i.e. changes in the pivotal for stress response and for the reproduction pituitary-adrenal and pituitary-gonadal systems.

The historical overview is presented of genetic experiments in L.V. Krushinsky’s laboratory in Moscow State University. L.V. Krushinsky stated the three-component concept of animal behavior. He claimed that animal behavior has not only innate species specific behavior and the learning ability, but should be supplemented by another mental category, reasoning the ability for elementary logic operarions. Being rather lonesome at the beginning, Krushinsky got the spiritual support from D.K. Belyaev and B.L. Astaurov. The attempt to study the genetic bases of reasoning ability was performed in Krushinsky’s lab using the trait “extrapolation problem solving”, which meant the ability of an unexperienced naïve animal to find the food bait when it moved aside and disappeared from (not “in”) the view. The selection for high scores of this trait in the hybrid rat population (Norway rat × laboratory strain cross) was started. Initially the hybrid rats solved this problem in the statistically significant proportions, while the animals from further selection generations demonstrated the dramatic increase of anxiety (in spite of extensive handling of these animals), which made further experiments impossible. Much later another selection experiment started in which mice of a genetically heterogeneous population were selected for high scores of extrapolation problem and concomitantly for the lack-ofanxiety signs during the testing procedure. This selection for a cognitive trait produced some positive results, although the direct response to selection was very weak. The data obtained show the intricate connection between the mouse ability to solve the problem and the processes of anxiety, which in turn looks as non-uniform by its nature and mechanisms. The data from experiments performed in classical genetics should be combined with the new knowledge concerning the role of single genes determining animal behavior.

The study of genetic mechanisms, which underlie normal and abnormal behavioral traits, are important not only for fundamental knowledge of CNS function, but also for human well-being, as well as in the aspect of treatment of brain diseases. Accumulation of knowledge concerning the functions of genes, which are expressed in the CNS and are involved in the specific brain functions determined now by the success of molecular genetics, but it could not overshadow the importance of phenotype expression investigation of genetically determined traits, especially pathological ones. Epilepsy, as one of CNS diseases, occupies an important place in the row of these studies. In spite of numerous anticonvulsant drugs a significant proportion of epilepsy cases are still resistant to drug treatments. It means that the study of various aspects of epileptogenesis using animal models should be welcomed as it will help to elucidate those aspects of increased seizure proneness that are now out of scope of research attention. The distinct trait of this domain, which is not easy to analyze genetically, is audiogenic epilepsy (the seizure attack in response to loud sound). The important feature of audiogenic epilepsy is that the intense tonic seizures could be induced and analyzed repeatedly which makes it possible to induce the seizures repeatedly in the course of chronical experiments with the same animal, which is not possible in the cases of pharmacologically or electrically induced seizures. The Krushinsky – Molodkina (KM) inbred rat strain, which was the first among strains selected for audiogenic epilepsy, is briefly characterized. The specific seizure pattern, the rodent proneness for audiogenic epilepsy, as well as abnormal postictal states (catalepsy, in particular) illustrate the importance of this phenomenon not only as a model of certain brain pathology, but also as a matter of general biological importance. The importance of genetic background factor (demonstrated in the case of study of comorbidity (audiogenic epilepsy, anxiety and depression)) could be crucial in the studies of other CNS anomalies. The audiogenic seizures as a physiological phenomenon which often accompanies several brain pathologies had been in the scope of research interests for D.K. Belyaev and his colleagues in his Institute.

Domesticated mammals of many different species share a set of physical and physiological traits that are not displayed by any of their wild progenitors. This suite of traits, now termed the “domestication syndrome” (DS), has been a puzzle since Charles Darwin discovered it. Two general explanations of its basis have been proposed, which in principle, could also apply to other vertebrates, such as fish and birds, whose domesticated varieties show some of its elements. The two ideas are termed here, respectively, the thyroid hormone hypothesis or the THH, and the neural crest cell hypothesis, the NCCH. The two ideas make distinctly different genetic predictions. Here, the current relevant evidence from genomics is evaluated and it is concluded that the NCCH has more support. Nevertheless, one set of observations, from chickens, suggest a potentially important role of altered thyroid metabolism in domestication. In addition, recent studies indicate the possibility of additional genetic factors in domestication, affecting tameness and sociality, that may go beyond either hypothesis. The tasks that lie ahead to fully ascertain the genetic bases of the “domestication syndrome” and the behaviors that characterize mammalian domestication are discussed briefly.

The domestication syndrome already recognized by Darwin shows that domesticated species acquire a number of novel morphological, physiological and behavioral characteristics not present in their wild ancestors. Because body size and sexual size dimorphism (SSD) are essential characteristics of species that affect most aspects of their life histories, we studied the effects of domestication on body size and SSD in domestic dogs and cats in comparison with their wild relatives: the Canidae and Felidae, respectively, and also analyzed the occurrence of Rensch’s rule within both domestic species. We studied maximum body mass and maximum height at withers of 64 and 89 domestic dog breeds respectively, and maximum body mass of 37 domestic cat breeds as well as body mass data for 36 wild Canidae and 36 wild Felidae from our previous studies. Our results have shown that domestic dogs maintain a level and range of body mass which largely exceeds that of the Canidae as a whole while maintaining a similar degree and range of SSD. On the contrary, domestic cats show a much reduced body mass range within the limits of their ancestor species while showing comparable levels of SSD as shown by the Felidae. Regarding Rensch’s rule, both Reduced Major Axis and Ordinary Least Squares regressions showed that both domestic species present a scaling of male and female body sizes consistent with Rensch’s rule while their wild relatives do not. We discuss these findings in the light of present knowledge about the domestication of Canis familiaris and Felis catus.

Geometric morphometrics is widely used to study underlying causal factors in phenotypic evolution and to reconstruct evolutionary history of phenotypes. However, non-linearities in the phenotypic landscape may exist such that analytical solutions derived from comparison of phenotypes in morphospace may have complex or contradictory relationships in the space of the underlying factors. Ancestral reconstruction of horn morphology based on two mammalian ungulates illustrates how biologically improbable results can arise from the mathematical properties of geometric morphometric morphospaces. Raup’s shell coiling equations are used to illustrate the potential for contradictory conclusions to be drawn from ancestral reconstructions in parameter spaces (such as measurements of levels of gene expression or allele frequencies) versus shape spaces (such as morphospaces based on phenotypic analysis). These examples are generalizable to many real morphometric studies, suggesting that care should be taken when drawing conclusions about genetic, developmental, or environmental processes based on morphometric analyses. Dense sampling of shape space and the use of fully multivariate and, perhaps, nonlinear methods can help forestall potential problems.

This paper commemotates centenary anniversary of Professor D.K. Belyaev and discusses his studies of photoperiodic changes on mammalian reproduction and embryonic development in the realm of modern data. Georgian white mutation (W G) in silver-black foxes was used as a model for such investigations. Ability of W G/W G homozygotes in standard photoperiodic conditions to go successfully through the implantation process is sharply reduced. Experimental elongation of daily light period for pregnant females from W G/w × W G/w crosses led to a very significant increase of W G/W G homozygotes in the progeny. Georgian white mutation likely causes a low expression of KIT gene. Presented here considerations of KIT protein and its ligand KITLG molecular and cellular interactions suggest a possibility for slow development of W G/W G blastocysts; hence many such blastocysts might be late and miss an opportunity for successful implantation. The discussed experimental photoperiodic change might influence a number of processes including developmental acceleration of W G/W G blastocysts, slow down trophopblast outgrowth of almost mature blastocysts, promotion of endometrial readiness and possible delay of the implantation start. All these processes could create more supportive conditions for successful implantation of W G/W G blastocysts.

In the paper the principles and methods of gene pools preservation of domestic animals breeds are explored. Currently 17 % of world species are under threat of extinction. The arguments in favor of preserving local breeds are: 1) economic factors, such as moderation of quality requirements for food products, moderation of hygienic and climatic conditions (new types of diseases, vaccines, changes in the external environment), necessity to keep a reserve for successful breeding, opportunity to use local breeds in harsh environmental conditions; 2) scientific factors, in particular, possibility of studying the gene pool of local breeds in order to reconstruct the evolution of domesticated species (exploring local breeds can reveal the mechanisms of the evolutionary processes, ontogeny, behavior patterns, natural and artificial selection); 3) cultural and historical factors related to the fact that native breeds represent living monuments of traditional culture of the population. Breeding is only possible while main-taining its inter- and interbreed genetic diversity, which can be identified through using various molecular genetic techniques. The examples of using new genetic technologies while preserving local breeds of two domesticated species are provided: sheep (Ovis aries) and cattle (Bos taurus). Using ISSR-fingerprinting (Inter Simple Sequence Repeats) some data were obtained, allowing us to determine the coefficient of genetic originality, to analyze population structure, to uncover similarities and differences, to identify and evaluate consolidation, breed purity and genealogical relationships of gene pools in a number of species (inbreeding). Within breeds of the same species or within one species it is possible to spot the ancient or the most estranged from the “protogene pool” groups of animals and, thus, to use these data in the breeding strategy aimed at the preservation of animal forms close to the ancestral form. Using various DNA markers, methods of genomic and genetic breeding in nature conservation and genetic selection of domesticated species provides an opportunity to obtain data on the genetic potential (value, originality) of the breed, which is important for scientific justification of its preservation.

Ever since Darwin published On The Origin of Species in 1859, the evolution of altruism has been a perennial paradox for evolutionary biologists. In this review, I will discuss three evolutionary paths to altruism – genetic relatedness, reciprocity, and group selection – and examine very recent work that uses social network modeling to help us better understand the evolution of altruism.

This article was inspired by some works of Belyaev in which he discussed the biosocial human nature, and is dedicated to the 100 year anniversary of Belyaev’ birth. The main views and theories of the human nature and the main paths of genome and culture evolution during anthropogenesis are overviewed. The views of the leading past and present world-wide scientists on the basic principles of the genome-culture interactions are discussed. The seeming contradictions between “geneticists” and “sociologists” are resolved due to the development of the concept of co-evolution of the genome and culture. The evolution of genes and culture is a whole, but not separate processes. During their evolution, humans modify and reconstruct their sociocultural environment. The new environment creates new selection vectors and forms new ways and directions for genetic evolution. In response to the culture development, the genetic determination of human cognitive processes also changed. A good example of the co-evolution of culture and the genome is the development of human speech, the voice and mimic apparatus of communication. The vital importance of the new possibilities for social communication in human populations disposes to the fixation of genetic features that facilitate such communication. At the early stages of human evolution, the areas of the cerebral cortex that were responsible for the development of speech expanded and reorganized. The point of genome-culture co-evolution is well illustrated by niche construction theory. It is emphasized that the evolution of culture, which is widely regarded as a non-biological phenomenon, is quite similar in all its respects to the Darwinian process, as a result of which a sociocultural environment that is adaptive for a given community is formed through selection and transmission in a series of generations. In fact, we are talking about epigenetic inheritance in the evolution of culture and morality. The review details the views and theoretical concepts of geneticists, sociologists, social psychologists, anthropologists, behaviorists and ethologists in explaining the phenomenon of human evolution. Some contradictions caused by different rates of evolution of culture and genome are considered. In conclusion, it is shown that Belyaev’s interpretation of the biosocial nature of humans finds confirmation in modern studies by geneticists, sociologists and psychologists.