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Evolution of the Avian Brain and Intelligence Current Biology Vol 15 No 23 R946 resources for the sea urchin can the structure of the avian brain be found at Primer relates to that of the mammalian http://www.ncbi.nlm.nih.gov/geno brain, and how names have had a me/guide/sea_urchin/ and at negative influence on how birds http://hgsc.bcm.tmc.edu/projects/ Evolution of the are perceived. seaurchin/. Negativity surrounding the avian avian brain and brain began in the late nineteenth Where can I find out more? intelligence century, when Ludwig Edinger Cameron, C.B., Garey, J.R., and Swalla, B.J. provided names for the various (2000). Evolution of the chordate body parts of the vertebrate brain. His plan: New insights from phylogenetic analyses of deuterostome phyla. Proc. Nathan J. Emery and form of nomenclature was based Natl. Acad. Sci. USA 97, 4469–4474. Nicola S. Clayton on the naïve view that evolution Davidson, E.H., Rast, J.P., Oliveri, P., occurs in a linear progression, so Ransick, A., Calestani, C., Yuh, C.H., In Western society, the term ‘bird that each new species is an Minokawa, T., Amore, G., Hinman, V., brain’ is often used as a elaboration of an older species. Arenas-Mena, C., et al. (2002). A genomic regulatory network for derogatory term for a person of This scala naturae is often development. Science 295, 1669–1678. diminished intellect, partly represented as a ladder. With Davidson, E.H., and Cameron, R.A. (2002). because many people tend to respect to intelligence, Arthur Arguments for sequencing the genome think of birds as pecking Jenson, one of the key figures in of the sea urchin, Strongylocentrotus machines, responding reflexively studies of human intelligence purpuratus. A white paper submitted to the National Human Genome Research to stimuli in their environment, and argued that “single-cell Institute. partly because birds seem so protozoans, such as amoeba, rank http://www.genome.gov/Pages/Researc different from us, with their beady at the bottom of the scale, h/Sequencing/SeqProposals/SeaU- eyes and small heads. But over 40 followed in order by the rchin_Genome.pdf Duboc, V., Rottinger, E., Lapraz, F., years ago William Thorpe, who invertebrates, the lower Besnardeau, L., and Lepage, T. (2005). was the leading authority on bird vertebrates, the lower mammals... Left-right asymmetry in the sea urchin learning at that time, pointed out: and finally the primates, in order: embryo is regulated by nodal signaling “The poor development in birds of New World monkeys, Old World on the right side. Dev. Cell 9, 147–158. any brain structures clearly monkeys, the apes, and at the Ettensohn, C.A., Wessel, G.M., and Wray, G.A. eds. (2004). Development of sea corresponding to the cerebral pinnacle, humans”. urchins, ascidians and other invertebrate cortex of mammals led to the With respect to brain evolution, deuterostomes: Experimental assumption among neurologists Edinger applied this scala naturae Approaches, Volume 74 (London: not only that birds are primarily suggesting that the brains of living Elsevier Academic Press). creatures of instinct, but also that vertebrates retained ancestral Hinman, V.F., Nguyen, A.T., Cameron, R.A., and Davidson, E.H. (2003). they are very little endowed with structures, but that new brain Developmental gene regulatory network the ability to learn...this areas were added onto older architecture across 500 million years of misconceived view of brain ones, or older areas increased in echinoderm evolution. Proc. Natl. Acad. mechanisms, hindered the size and complexity to form new Sci. USA 100, 13356–13361. Peterson, K.J., Arenas-Mena, C., and development of experimental areas (Figure 1A). According to Davidson, E.H. (2000). The A/P axis in studies on bird learning”. this view, evolutionarily older echinoderm ontogeny and evolution: In the 1960s little was known brains are simple, and so produce evidence from fossils and molecules. about the cognitive capacities of simple instinctive behaviour, and Evol. Dev. 2, 93–101. birds, but recent studies lend evolutionarily newer brains are Raff, E.C., Popodi, E.M., Kauffman, J.S., Sly, B.J., Turner, F.R., Morris, V.B., and Raff, support for Thorpe’s view: we now complex, and therefore can R.A. (2003). Regulatory punctuated know that some bird species control learned and intelligent equilibrium and convergence in the make and use tools, can count, behaviour. The oldest brain evolution of developmental pathways in remember specific past events regions — those present in all direct-developing sea urchins. Evol. Dev. and reason about the mental vertebrates — were prefixed with 5, 478–493. Shu, D.-G., Conway Morris, S., Han, J., states of individuals, behaviours the term ‘paleo-‘, the next oldest Zhang, Z.-F., and Liu, J.-N. (2004). that some have considered to be brain regions were given the prefix Ancestral echinoderms from the unique to humans. Despite the ‘archi-’, whereas the new brain Chengjiang deposits of China. Nature apparent cognitive similarity regions — those present in the 430, 422–428. between humans and some birds, species closest to the top of the neuroscientists have tended to ‘ladder’ — were assigned the 1 Molecular Genetics Program, Benaroya view bird brains as interesting prefix ‘neo-’. Research Institute at Virginia Mason, 1201 Ninth Avenue, Seattle, WA 98101, curiosities with little relevance to We now know that, as with USA. 2Department of Biology, University the workings of the human brain. other parts of the body, the brains of Washington, 24 Kincaid Hall, Seattle, Recently, however, the Avian of distantly related species tend to Washington 98195, USA. 3Department Brain Nomenclature Consortium be derived from the same basic of Molecular & Cellular Biology, published a series of papers elements found in the common Sunnybrook & Women’s Research attempting to re-address the issue ancestor — they exhibit homology Institute and Medical Biophysics, University of Toronto, 2075 Bayview of the importance of the bird brain (Figure 1B). So although the Avenue, Toronto, Ontario M4N 3MS, to neuroscience by investigating common ancestor of birds and Canada. how the avian brain evolved, how mammals lived approximately 300 Magazine R947 million years ago, studies of A B extant reptiles have revealed that Humans Complex Humans the reptilian (therapsid and sauropsid) forebrain is pallial in Non-primate Mammals origin and so the common Birds mammals ancestor should also have shared Reptiles this trait. If so, then the forebrain Birds of modern birds and mammals will Amphibians also be pallial. This seems to be Reptiles the case. Fish Edinger mistakenly thought that Amphibians the majority of the avian brain was Fish derived from the striatum or basal Simple Current Biology ganglia, which tends to be involved in species-specific Figure 1. Schematic representation of two theories of brain evolution. instinctual behaviours, such as (A) The outdated ‘scala naturae’ theory, where evolution occurs in a linear, progressive feeding and sexual and parental fashion up a ladder in which ‘lower’ (simple) species evolve into ‘higher’ (complex) behaviour, as well as responses to species; going from fish and amphibians at the bottom through reptiles and birds to primates and humans at the top. With respect to brain evolution, the increasing rewards and motor coordination. complexity resulting from climbing the ladder leads to the appearance of completely In mammals, the basal ganglia has new areas which are then added onto old ones. Each colour represents a different a striated appearance due to the hypothetical brain region, either old or new. (B) The modern theory, where evolution is fibre bundles running through it. tree-like and new species evolve from older ancestral forms. With respect to brain Areas in the avian forebrain evolution, complexity is derived from refining neural structures which are already tended to resemble the present in ancestral forms, such that brain regions increase in size. There are no truly new brain areas, only elaborations of established regions. The colours represent mammalian striatum and so most different brain regions, but rather than new areas being added, evolutionarily old areas areas (paleo-, archi- or neo-) were are increased or decreased in size (or complexity). named with the root word ‘striatum’. For example, the avian matter. The analogy would be to intelligent. The names have cerebrum was originally separated compare a club sandwich changed, but not necessarily the into neostriatum, hyperstriatum, (mammalian) to a pepperoni pizza abilities associated with them. archistriatum, paleostriatum and (avian). There are over 9,000 species of ectostriatum (Figure 2). Note that The accumulation of new data birds, however many of those very few structures were thought led a consortium of avian studied by comparative to be derived from the pallium, neuroscientists to propose a new psychologists are not well and named with the root words nomenclature for the avian brain, endowed with the cognitive skills ’cortex’ or ‘pallium’. By contrast, where the forebrain was renamed associated with primates and much of the mammalian forebrain in the light of this modern dolphins. The humble pigeon, for was known to be derived from the understanding of its evolution and example, is a master at visual pallium, which resulted in a six- the relative contributions of the discrimination and memory: it can layered neocortex, the area of the striatum and pallium (Figure 2). remember hundreds of different brain which is involved in thinking, The proportion of the forebrain objects for long periods, reasoning and planning. that is known to be cortical-like discriminate between different Research in the last 30 years on (pallial) in structure has increased painting styles that would stump neural connectivity, gene significantly (Figure 2). These art history undergraduates, and expression and lesions has name changes reflect our current navigate hundreds of miles by caused a revision in our thinking understanding of how these brain following the trajectory of major about the avian forebrain and the regions evolved (Table 1), but they roads.
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