Supernumerary scutes verify a segment-dependent model of the horny shell development in turtles.

Turtle horny shell has a scute pattern, which is conservative through evolution and across species. The discovery of epidermal placodes as the scute primordia and their strict topographical association to the somites of the turtle embryo suggested a new interpretation of the developmental mechanism of the scute pattern. Here, we tested the hypothesis that horny scutes develop from a mosaic of placodes corresponding exactly to the paths of myoseptae, with vertebral and pleural scutes developing staggered in adjacent segments, and marginal scutes developing in every segment. This scheme predicts little variation in marginals and suggests intercalary supernumerary scutes as potential variations for the vertebral and pleural rows. We examined spatial and numerical variations of the horny shell in 655 newly hatched olive ridley sea turtle, Lepidochelys olivacea, which is known to have a highly variable horny shell. In total, 120 patterns of carapacial scutes and 10 patterns of scutes on plastron, differing in the number and position of scutes were found. The number of vertebral scutes varied from 4 to 10. Variations with five, six and seven vertebrals occurred with the greatest and nearly equally frequency (31.5% on average). Pleural scutes were from 5 to 10 at one or both sides, and the typical symmetric pattern for sea turtles with five pairs of pleurals was only seen in ca. 12% of specimens. In contrast, the majority of the specimens (92.7%) had just 13 pairs of marginals, showing a stable normal pattern. Similarly, on plastron the horny scutes were conservative, too; about 85% of specimens standardly had six pairs of plastral scutes and all specimens had four pairs of inframarginals. Despite a high level of variation of vertebral and pleural scutes in olive ridley turtle, all patterns fall into the theoretical spectrum of possible variants predicted by the segment-dependent model of development of the turtle horny shell. Therefore, the results of our analysis support the existence of direct morphogenetic correlation between the number and distribution of normal and supernumerary scutes and metamere organization of the turtle embryo.

What do wild saiga antelopes tell us about the relative roles of the two brain hemispheres in social interactions?

Two brain hemispheres are unequally involved in the processing of social stimuli, as demonstrated in a wide range of vertebrates. A considerable number of studies have shown the right hemisphere advantage for social processing. At the same time, an approach-withdrawal hypothesis, mainly based on experimental evidence, proposes the involvement of both brain hemispheres according to approach and withdrawal motivation. The present study aimed to test the relative roles of the two hemispheres in social responses displayed in a natural context. Visual biases, implicating hemispheric lateralization, were estimated in the social interactions of saiga antelope in the wild. In individually identified males, the left/right visual field use during approach and withdrawal responses was recorded based on the lateral head/body position, relative to the conspecific. Lateralized approach responses were investigated in three types of interactions, with left visual field bias found for chasing a rival, no bias-for attacking a rival, and right visual field bias-for pursuing a female. In two types of withdrawal responses, left visual field bias was found for retreating after fighting, while no bias was evident in fight rejecting. These findings demonstrate that neither the right hemisphere advantage nor the approach-withdrawal distinction can fully explain the patterns of lateralization observed in social behaviour. It is clear that both brain hemispheres play significant roles in social responses, while their relative contribution is likely determined by a complex set of motivational and emotional factors rather than a simple dichotomous distinction such as, for example, approach versus withdrawal motivation.

Facing each other: mammal mothers and infants prefer the position favouring right hemisphere processing.

The right hemisphere plays a crucial role in social processing. Human mothers show a robust left cradling/holding bias providing greater right-hemispheric involvement in the exchange of social information between mother and infant. Here, we demonstrate that a similar bias is evident in face-to-face spatial interactions in marine and terrestrial non-primate mammals. Walruses and Indian flying foxes showed a significant population-level preference for the position which facilitates the use of the left visual field in both mother and infant. This behavioural lateralization may have emerged owing to benefits conferred by the enhanced right-hemispheric social processing providing the mother and infant an optimal perception of each other.

Lateralization of mother-infant interactions in wild horses.

The manifestation of behavioural lateralization has been shown to be modified by environmental conditions, life experiences, and selective breeding. This study tests whether the lateralization recently found in feral domestic horse (Equus caballus) is evident in undomesticated horses. Mother-offspring interactions were investigated in Przewalski's horse (E. ferus przewalskii) living in their natural habitat in Mongolia. Lateral position preferences during mare-foal spontaneous reunions were used as a behavioural marker of visual lateralization. Preferences were separately assessed for foals' approaches to their mothers and mares' approaches to their foals. Preference to keep the mother in the visual field of the left eye was found in various types of foals' behaviour. In slow travelling, Przewalski's foals showed stronger preference for the left eye use than feral horse foals. Population-level left-eye bias was also found in mothers approaching their foals. Our results indicate right-hemispheric dominance for control of mother-offspring interactions in Przewalski's horses, similar to what has been reported for other mammals including humans. Benefits conferred by the lateralized social processing of and responding to social stimuli may explain that the left-lateralized social behaviour is a robust trait of equine behaviour, not modified by domestication or specific environmental conditions of the population.

Lateralization of mother-infant interactions in a diverse range of mammal species.

Left-cradling bias is a distinctive feature of maternal behaviour in humans and great apes, but its evolutionary origin remains unknown. In 11 species of marine and terrestrial mammal, we demonstrate consistent patterns of lateralization in mother-infant interactions, indicating right hemisphere dominance for social processing. In providing clear evidence that lateralized positioning is beneficial in mother-infant interactions, our results illustrate a significant impact of lateralization on individual fitness.

Early expression of manual lateralization in bipedal marsupials.

Robust lateralization in forelimb use has recently been found in bipedal, but not quadrupedal, marsupial mammals. The link between bipedality and handedness, occurring in both marsupials and primates, remains to be investigated. To shed light on the developmental origins of marsupial manual lateralization, infants of macropod marsupials were examined before and shortly after the acquisition of habitual bipedal posture and locomotion. Forelimb preferences were assessed in natural, not artificially evoked, behaviors of infant red-necked wallaby in the wild and infant eastern gray kangaroo in free-ranging captivity. Pouch young of both species showed population-level left-forelimb preference when manipulating food objects, such as leaves and grass blades. This result provides the first report of lateralization in pouch young marsupials and rare evidence of lateralized manual activity in early mammalian ontogenesis. Young-at-foot juveniles of eastern gray kangaroo preferred to use the left forelimb to manipulate the mother's pouch edge as previously shown for red-necked wallaby. In both species, the direction of biases in manipulative behavior for young-at-foot and pouch young was the same as in adults. Forelimb preferences in offspring were positively correlated with the forelimb preferences of their mothers. Our results strongly suggest that the emergence of individual and population-level forelimb preferences in macropod infants precedes the onset of independent standing and locomotion. In all probability, manual lateralization in bipedal marsupials, such as kangaroos and wallabies, is not determined by the acquisition of habitual bipedality in the course of ontogenesis. (PsycINFO Database Record

Primary Cilia as a Possible Link between Left-Right Asymmetry and Neurodevelopmental Diseases.

Cilia have multiple functions in the development of the entire organism, and participate in the development and functioning of the central nervous system. In the last decade, studies have shown that they are implicated in the development of the visceral left-right asymmetry in different vertebrates. At the same time, some neuropsychiatric disorders, such as schizophrenia, autism, bipolar disorder, and dyslexia, are known to be associated with lateralization failure. In this review, we consider possible links in the mechanisms of determination of visceral asymmetry and brain lateralization, through cilia. We review the functions of seven genes associated with both cilia, and with neurodevelopmental diseases, keeping in mind their possible role in the establishment of the left-right brain asymmetry.

Asymmetry of righting reflexes in sea turtles and its behavioral correlates.

The righting responses, when the animal rights itself over one side of the body after been overturned on the back, are one of the simplest ways to test for laterality, especially in lower vertebrates. In anuran amphibians unilateral preferences in righting responses correlated to the degree of the use of alternating-limb (asynchronous) movements during locomotion. Turtles is one of the underrepresented vertebrate groups in the studies of laterality, while possess also different types of locomotion (with synchronous or asynchronous use of the contralateral limbs), which allows testing the hypothesis on functional relationship between the mode of locomotion and the strength of laterality. We studied two species of sea turtles, Green turtle (Chelonia mydas) and Olive Ridley turtle (Lepidochelys olivacea), which differ from the majority of other representatives of the order in that they mostly utilize synchronous locomotion, when all four limbs move simultaneously in strokes (scratching). In righting response tests turtles demonstrated individual and weak population level laterality, which differed in strength. The Green turtle was less lateralized with the majority of individuals being ambipreferent. The Olive Ridley turtle had a greater number of lateralized individuals and a greater average strength of laterality. Interspecies comparison to land tortoises, which use only asynchronous (alternating-limb) walking (crawling), confirmed the rule found in amphibians: the more asynchronous locomotion is used, the greater is the strength of laterality in righting. Hence, data from turtles and amphibians may represent a phenomenon common for all quadruped vertebrates. We also discuss possible biomechanical and neurological correlates of this evolutionary change in locomotory patterns and lateralization in sea turtles when adapting to sea life.

Evidence for the perceptual origin of right-sided feeding biases in cetaceans.

Foraging behaviour of many cetacean species features the side biases at the population level. The origin of these behavioural lateralisations remains generally unclear. Here we explored lateralisation in aerial display of resident orcas in different behavioural contexts. Side preferences were analysed in lunging during foraging and breaching. One event of each type of displays per individually identified orca was used for analysis. Orcas showed a population-level preference to lunge on the right side when foraging (75% of lunges). In contrast, no lateralisation was found in breaching (54% of breaches to the right, 45% to the left). The right-sided bias in foraging found in orcas is in line with evidence from other whales, both baleen and toothed, and confirms the uniformity of feeding biases among cetaceans. In contrast to breaching, lunging in orcas was associated with fish pursuit, that is, with focused attention to and sensory perception of prey stimulus. The emergence of lateralisation in lunging and the absence of significant bias in breaching suggest that feeding biases in whales are underpinned by sensory lateralisation, that is, by lateralised hemispheric processing of sensory information about the prey. Evidence from orcas may be extrapolated to other cetaceans since right-sided biases in lunging during foraging is a very widespread phenomenon and likely to have a common origin. Our findings support the hypothesis that right-sided feeding biases are determined by left-hemisphere specialisation.

Parallel Emergence of True Handedness in the Evolution of Marsupials and Placentals.

Recent studies have demonstrated a close resemblance between some handedness patterns in great apes and humans. Despite this, comparative systematic investigations of manual lateralization in non-primate mammals are very limited. Among mammals, robust population-level handedness is still considered to be a distinctive human trait. Nevertheless, the comprehensive understanding of handedness evolution in mammals cannot be achieved without considering the other large mammalian lineage, marsupials. This study was designed to investigate manual lateralization in non-primate mammals using the methodological approach applied in primate studies. Here we show that bipedal macropod marsupials display left-forelimb preference at the population level in a variety of behaviors in the wild. In eastern gray and red kangaroos, we found consistent manual lateralization across multiple behaviors. This result challenges the notion that in mammals the emergence of strong "true" handedness is a unique feature of primate evolution. The robust lateralization in bipedal marsupials stands in contrast to the relatively weak forelimb preferences in marsupial quadrupeds, emphasizing the role of postural characteristics in the evolution of manual lateralization as previously suggested for primates. Comparison of forelimb preferences in seven marsupial species leads to the conclusion that the interspecies differences in manual lateralization cannot be explained by phylogenetic relations, but rather are shaped by ecological adaptations. Species' postural characteristics, especially bipedality, are argued to be instrumental in the origin of handedness in mammals.

Beta-catenin in schizophrenia: Possibly deleterious novel mutation.

Schizophrenia is a debilitating psychiatric disorder, affecting approximately 1% of the human population. Mostly genetic factors contribute to schizophrenia, but the genetics are complex and various aspects of brain functioning and structure, from development to synapse plasticity, seem to be involved in the pathogenesis. The goal of the study was to look for novel mutations in genes, implicated in molecular networks, important in schizophrenia. In the study four candidate genes taking part in the WNT signaling pathway were analyzed by sequencing in a cohort of 87 schizophrenia patients from Saint Petersburg, Russia. The gene list included CTNNB1 (beta-catenin), GSK3B, WNT2B and WNT7B. The impact of discovered variants on the protein function was analyzed in silico. We found three variants in the genes CTNNB1 and WNT7B, absent in healthy controls, including 212 controls from the same geographic area. The novel mutation c.1943A>G (p.N648S) in CTNNB1 seems to be the best candidate for disease-associated mutation in this study, as it damages the protein product in silico. This is the first study reporting mutations in CTNNB1 in schizophrenia.

Sequencing of five left-right cerebral asymmetry genes in a cohort of schizophrenia and schizotypal disorder patients from Russia.

OBJECTIVE: Schizophrenia is a severe psychiatric disorder, affecting ∼1% of the human population. The genetic contribution to schizophrenia is significant, but the genetics are complex and many aspects of brain functioning, from neural development to synapse structure, seem to be involved in the pathogenesis. A novel way to study the molecular causes of schizophrenia is to study the genetics of left-right (LR) brain asymmetry, the disease feature often observed in schizophrenic patients. METHODS: In this study, we analyzed by sequencing five candidate LR cerebral asymmetry genes in a cohort of 95 schizophrenia and schizotypal disorder patients from Saint Petersburg, Russia. The gene list included LMO4, LRRTM1, FOXP2, the PCDH11X/Y gene pair, and SRY. RESULTS: We found 17 previously unreported variants in the genes LRRTM1, FOXP2, LMO4, and PCDH11X in the 3'-UTR and 5'-UTR. The variants might contribute toward an altered mRNA processing, which could lead to altered mRNA amounts in developing neurons of the brain and establishment of an incorrect LR asymmetry profile. CONCLUSION: This is the first study in which multiple candidate genes for cerebral LR asymmetry and schizophrenia have been analyzed by sequencing. The approach to study the genetics of schizophrenia from the perspective of an LR cerebral asymmetry disturbance deserves more attention.

Lateralisation of rotational swimming but not fast escape response in the juvenile sterlet sturgeon, Acipenser ruthenus (Chondrostei: Acipenseridae).

For the first time, behavioural lateralisation was shown in a chondrostean fish (sterlet sturgeon Acipenser ruthenus). A significant directional bias was found in young A. ruthenus swimming along a circular swimway. This laterality manifested itself as an individual preference for a certain movement direction (either clockwise or counterclockwise) which was consistent at the retest 10 days later. On the other hand, no significant rotational bias was observed at the population level. The same sterlet individuals displayed the C-start (the first stage of escape response) elicited by sudden low-frequency sound vibrations (50 Hz). However, the experiments failed to reveal either individual or population laterality of this response: the frequencies of leftward and rightward bends in startled fish were virtually equal. These results demonstrate that the two types of laterality can be independent in fish.

Forelimb preferences in quadrupedal marsupials and their implications for laterality evolution in mammals.

BACKGROUND: Acquisition of upright posture in evolution has been argued to facilitate manual laterality in primates. Owing to the high variety of postural habits marsupials can serve as a suitable model to test whether the species-typical body posture shapes forelimb preferences in non-primates or this phenomenon emerged only in the course of primate evolution. In the present study we aimed to explore manual laterality in marsupial quadrupeds and compare them with the results in the previously studied bipedal species. Forelimb preferences were assessed in captive grey short-tailed opossum (Monodelphis domestica) and sugar glider (Petaurus breviceps) in four different types of unimanual behaviour per species, which was not artificially evoked. We examined the possible effects of sex, age and task, because these factors have been reported to affect motor laterality in placental mammals. RESULTS: In both species the direction of forelimb preferences was strongly sex-related. Male grey short-tailed opossums showed right-forelimb preference in most of the observed unimanual behaviours, while male sugar gliders displayed only a slight, not significant rightward tendency. In contrast, females in both species exhibited consistent group-level preference of the left forelimb. We failed to reveal significant differences in manual preferences between tasks of potentially differing complexity: reaching a stable food item and catching live insects, as well as between the body support and food manipulation. No influence of subjects' age on limb preferences was found. CONCLUSIONS: The direction of sex-related differences in the manual preferences found in quadrupedal marsupials seems to be not typical for placental mammals. We suggest that the alternative way of interhemispheric connection in absence of corpus callosum may result in a fundamentally distinct mechanism of sex effect on limb preferences in marsupials compared to placentals. Our data confirm the idea that non-primate mammals differ from primates in sensitivity to task complexity. Comparison of marsupial species studied to date indicate that the vertical body orientation and the bipedalism favor the expression of individual- and population-level forelimb preferences in marsupials much like it does in primates. Our findings give the first evidence for the effect of species-typical posture on the manual laterality in non-primate mammals.

Eye as a key element of conspecific image eliciting lateralized response in fish.

Visual lateralization in different aspects of social behaviour has been found for numerous species of vertebrates ranging from fish to mammals. For inspection of a shoal mate, many fishes show a left eye-right hemisphere preference. Here, we tested the hypothesis that in fish, there is a key cue in the conspecific appearance, which elicits lateralized response to the whole image of the conspecific. In a series of eight experiments, we explored eye preferences in cryptic-coloured Amur sleeper, Perccottus glenii, fry. Fish displayed left-eye preferences at the population level for inspection of a group of conspecifics, their own mirror image, and a motionless flat model of a conspecific. In contrast, no population bias was found for scrutinizing an empty environment or a moving cylinder. When fry were showed a model of a conspecific in a lateral view with the eye displaced from the head to the tail, they again showed a significant preference for left-eye use. On the other hand, 'eyeless' conspecific model elicited no lateralized viewing in fry. Finally, the left-eye preference was revealed for scrutiny of the image of a conspecific eye alone. We argue that in Amur sleeper fry, eye is the element of the conspecific image, which can serve as a 'key' for the initiation of lateralized social response. This key element may serve as a trigger for the rapid recognition of conspecifics in the left eye-right hemisphere system. Possible causes and advantages of lateralized perception of social stimuli and their key elements are discussed in the context of current theories of brain lateralization.

Dual embryonic origin of the hyobranchial apparatus in the Mexican axolotl (Ambystoma mexicanum).

Traditionally, the cartilaginous viscerocranium of vertebrates is considered as neural crest (NC)-derived. Morphological work carried out on amphibian embryos in the first half of the XX century suggested potentially mesodermal origin for some hyobranchial elements. Since then, the embryonic sources of the hyobranchial apparatus in amphibians has not been investigated due to lack of an appropriate long-term labelling system. We performed homotopic transplantations of neural folds along with the majority of cells of the presumptive NC, and/or fragments of the head lateral plate mesoderm (LPM) from transgenic GFP+ into white embryos. In these experiments, the NC-derived GFP+ cells contributed to all hyobranchial elements, except for basibranchial 2, whereas the grafting of GFP+ head mesoderm led to a reverse labelling result. The grafting of only the most ventral part of the head LPM resulted in marking of the basibranchial 2 and the heart myocardium, implying their origin from a common mesodermal region. This is the first evidence of contribution of LPM of the head to cranial elements in any vertebrate. If compared to fish, birds, and mammals, in which all branchial skeletal elements are NC-derived, the axolotl (probably this is true for all amphibians) demonstrates an evolutionary deviation, in which the head LPM replaces NC cells in a hyobranchial element. This implies that cells of different embryonic origin may have the same developmental program, leading to the formation of identical (homologous) elements of the skeleton.

Does bipedality predict the group-level manual laterality in mammals?

BACKGROUND: Factors determining patterns of laterality manifestation in mammals remain unclear. In primates, the upright posture favours the expression of manual laterality across species, but may have little influence within a species. Whether the bipedalism acts the same in non-primate mammals is unknown. Our recent findings in bipedal and quadrupedal marsupials suggested that differences in laterality pattern, as well as emergence of manual specialization in evolution might depend on species-specific body posture. Here, we evaluated the hypothesis that the postural characteristics are the key variable shaping the manual laterality expression across mammalian species. METHODOLOGY/PRINCIPAL FINDINGS: We studied forelimb preferences in a most bipedal marsupial, brush-tailed bettong, Bettongia penicillata in four different types of unimanual behavior. The significant left-forelimb preference at the group level was found in all behaviours studied. In unimanual feeding on non-living food, catching live prey and nest-material collecting, all or most subjects were lateralized, and among lateralized bettongs a significant majority displayed left-forelimb bias. Only in unimanual supporting of the body in the tripedal stance the distribution of lateralized and non-lateralized individuals did not differ from chance. Individual preferences were consistent across all types of behaviour. The direction or the strength of forelimb preferences were not affected by the animals' sex. CONCLUSIONS/SIGNIFICANCE: Our findings support the hypothesis that the expression of manual laterality depends on the species-typical postural habit. The interspecies comparison illustrates that in marsupials the increase of bipedality corresponds with the increase of the degree of group-level forelimb preference in a species. Thus, bipedalism can predict pronounced manual laterality at both intra- and interspecific levels in mammals. We also conclude that quadrupedal position in biped species can slightly hinder the expression of manual laterality, but the evoked biped position in quadrupedal species does not necessarily lead to the enhanced manifestation of manual laterality.

Neural crest does not contribute to the neck and shoulder in the axolotl (Ambystoma mexicanum).

BACKGROUND: A major step during the evolution of tetrapods was their transition from water to land. This process involved the reduction or complete loss of the dermal bones that made up connections to the skull and a concomitant enlargement of the endochondral shoulder girdle. In the mouse the latter is derived from three separate embryonic sources: lateral plate mesoderm, somites, and neural crest. The neural crest was suggested to sustain the muscle attachments. How this complex composition of the endochondral shoulder girdle arose during evolution and whether it is shared by all tetrapods is unknown. Salamanders that lack dermal bone within their shoulder girdle were of special interest for a possible contribution of the neural crest to the endochondral elements and muscle attachment sites, and we therefore studied them in this context. RESULTS: We grafted neural crest from GFP+ fluorescent transgenic axolotl (Ambystoma mexicanum) donor embryos into white (d/d) axolotl hosts and followed the presence of neural crest cells within the cartilage of the shoulder girdle and the connective tissue of muscle attachment sites of the neck-shoulder region. Strikingly, neural crest cells did not contribute to any part of the endochondral shoulder girdle or to the connective tissue at muscle attachment sites in axolotl. CONCLUSIONS: Our results in axolotl suggest that neural crest does not serve a general function in vertebrate shoulder muscle attachment sites as predicted by the "muscle scaffold theory," and that it is not necessary to maintain connectivity of the endochondral shoulder girdle to the skull. Our data support the possibility that the contribution of the neural crest to the endochondral shoulder girdle, which is observed in the mouse, arose de novo in mammals as a developmental basis for their skeletal synapomorphies. This further supports the hypothesis of an increased neural crest diversification during vertebrate evolution.

Visual laterality of calf-mother interactions in wild whales.

BACKGROUND: Behavioral laterality is known for a variety of vertebrate and invertebrate animals. Laterality in social interactions has been described for a wide range of species including humans. Although evidence and theoretical predictions indicate that in social species the degree of population level laterality is greater than in solitary ones, the origin of these unilateral biases is not fully understood. It is especially poorly studied in the wild animals. Little is known about the role, which laterality in social interactions plays in natural populations. A number of brain characteristics make cetaceans most suitable for investigation of lateralization in social contacts. METHODOLOGY/PRINCIPAL FINDINGS: Observations were made on wild beluga whales (Delphinapterus leucas) in the greatest breeding aggregation in the White Sea. Here we show that young calves (in 29 individually identified and in over a hundred of individually not recognized mother-calf pairs) swim and rest significantly longer on a mother's right side. Further observations along with the data from other cetaceans indicate that found laterality is a result of the calves' preference to observe their mothers with the left eye, i.e., to analyze the information on a socially significant object in the right brain hemisphere. CONCLUSIONS/SIGNIFICANCE: Data from our and previous work on cetacean laterality suggest that basic brain lateralizations are expressed in the same way in cetaceans and other vertebrates. While the information on social partners and novel objects is analyzed in the right brain hemisphere, the control of feeding behavior is performed by the left brain hemisphere. Continuous unilateral visual contacts of calves to mothers with the left eye may influence social development of the young by activation of the contralateral (right) brain hemisphere, indicating a possible mechanism on how behavioral lateralization may influence species life and welfare. This hypothesis is supported by evidence from other vertebrates.

An eye for a worm: lateralisation of feeding behaviour in aquatic anamniotes.

Some animals, notably birds, preferentially approach and capture food items in their right visual field. However, this lateralised behaviour has not been studied extensively in anamniotes. Here we test eye preference during feeding for a fish, (Perccottus glenii; Teleostei, Perciformes), a newt, (Pleurodeles walti; Amphibia, Caudata), and a frog, (Xenopus laevis; Amphibia, Anura) using a test chamber that assesses reaction to visual stimuli while blocking olfactory and mechanical input. Both the fish and the newt showed right preferences in reactions to food items, but the frog did not. Our data extend our knowledge of the lateralised behaviours of vertebrates and are the first record of lateralised prey capture in a caudate amphibian. This finding dates back the history of the common pattern for visual lateralisation in vertebrates to Devonian, when the fish and quadruped lineages diverged.