The function(s) of consciousness: an evolutionary perspective.

The functions of consciousness, viewed from an evolutionary standpoint, can be categorized as being either general or particular. There are two general functions, meaning those that do not depend on the particulars of how consciousness influences behavior or how and why it first evolved: of (1) expanding the behavioral repertoire of the individual through the gradual accumulation of neurocircuitry innovations incorporating consciousness that would not exist without it, and (2) reducing the time scale over which preprogrammed behaviors can be altered, from evolutionary time, across generations, to real-time. But neither answers Velmans' question, of why consciousness is adaptive in a proximate sense, and hence why it would have evolved, which depends on identifying the particular function it first performed.

Mental causation: an evolutionary perspective.

The relationship between consciousness and individual agency is examined from a bottom-up evolutionary perspective, an approach somewhat different from other ways of dealing with the issue, but one relevant to the question of animal consciousness. Two ways are identified that would decouple the two, allowing consciousness of a limited kind to exist without agency: (1) reflex pathways that incorporate conscious sensations as an intrinsic component (InCs), and (2) reflexes that are consciously conditioned and dependent on synaptic plasticity but not memory (CCRs). Whether InCs and CCRs exist as more than hypothetical constructs is not clear, and InCs are in any case limited to theories where consciousness depends directly on EM field-based effects.

The Cambrian fossil Pikaia, and the origin of chordate somites.

The Middle Cambrian fossil Pikaia has a regular series of vertical bands that, assuming chordate affinities, can be interpreted as septa positioned between serial myotomes. Whether Pikaia has a notochord and nerve cord is less certain, as the dorsal organ, which has no obvious counterpart in living chordates, is the only clearly defined axial structure extending the length of the body. Without a notochord to serve as a reference point, the location of the nerve cord is then conjectural, which begs the question of how a dorsal neural center devoted to somite innervation would first have arisen from a more diffuse ancestral plexus of intraepithelial nerves.

Consciousness and its hard problems: separating the ontological from the evolutionary.

Few of the many theories devised to account for consciousness are explicit about the role they ascribe to evolution, and a significant fraction, by their silence on the subject, treat evolutionary processes as being, in effect, irrelevant. This is a problem for biological realists trying to assess the applicability of competing theories of consciousness to taxa other than our own, and across evolutionary time. Here, as an aid to investigating such questions, a consciousness "machine" is employed as conceptual device for thinking about the different ways ontology and evolution contribute to the emergence of a consciousness composed of distinguishable contents.

On the origins and evolution of qualia: An experience-space perspective.

This paper elaborates on a proposal for mapping a configuration space for selector circuits (SCs), defined as the subset of neural correlates of consciousness (NCCs) responsible for evoking particular qualia, to its experiential counterpart, experience-space (E-space), as part of an investigation into the nature of conscious experience as it first emerged in evolution. The dimensionality of E-space, meaning the degrees of freedom required to specify the properties of related sets of qualia, is at least two, but the utility of E-space as a hypothetical construct is much enhanced by assuming it is a large dimensional space, with at least several times as many dimensions as there are categories of qualia to occupy them. Phenomenal consciousness can then be represented as having originated as one or more multidimensional ur-experiences that combined multiple forms of experience together.

Patterning, From Conifers to Consciousness: Turing's Theory and Order From Fluctuations.

This is a brief account of Turing's ideas on biological pattern and the events that led to their wider acceptance by biologists as a valid way to investigate developmental pattern, and of the value of theory more generally in biology. Periodic patterns have played a key role in this process, especially 2D arrays of oriented stripes, which proved a disappointment in theoretical terms in the case of segmentation, but a boost to theory as applied to skin patterns in fish and model chemical reactions. The concept of "order from fluctuations" is a key component of Turing's theory, wherein pattern arises by selective amplification of spatial components concealed in the random disorder of molecular and/or cellular processes.

An evolutionary perspective on chordate brain organization and function: insights from amphioxus, and the problem of sentience.

The similarities between amphioxus and vertebrate brains, in their regional subdivision, cell types and circuitry, make the former a useful benchmark for understanding the evolutionary innovations that shaped the latter. Locomotory control systems were already well developed in basal chordates, with the ventral neuropile of the dien-mesencephalon serving to set levels of activity and initiate locomotory actions. A chief deficit in amphioxus is the absence of complex vertebrate-type sense organs.

Consciousness as a Product of Evolution: Contents, Selector Circuits, and Trajectories in Experience Space.

Conscious experience can be treated as a complex unified whole, but to do so is problematic from an evolutionary perspective if, like other products of evolution, consciousness had simple beginnings, and achieved complexity only secondarily over an extended period of time as new categories of subjective experience were added and refined. The premise here is twofold, first that these simple beginnings can be investigated regardless of whether the ultimate source of subjective experience is known or understood, and second, that of the contents known to us, the most accessible for investigation will be those that are, or appear, most fundamental, in the sense that they resist further deconstruction or analysis. This would include qualia as they are usually defined, but excludes more complex experiences (here, formats) that are structured, or depend on algorithmic processes and/or memory.

Amphioxus neuroglia: Molecular characterization and evidence for early compartmentalization of the developing nerve cord.

Glial cells play important roles in the development and homeostasis of metazoan nervous systems. However, while their involvement in the development and function in the central nervous system (CNS) of vertebrates is increasingly well understood, much less is known about invertebrate glia and the evolutionary history of glial cells more generally. An investigation into amphioxus glia is therefore timely, as this organism is the best living proxy for the last common ancestor of all chordates, and hence provides a window into the role of glial cell development and function at the transition of invertebrates and vertebrates.

Evolving Consciousness: Insights From Turing, and the Shaping of Experience.

A number of conceptual difficulties arise when considering the evolutionary origin of consciousness from the pre-conscious condition. There are parallels here with biological pattern formation, where, according to Alan Turing's original formulation of the problem, the statistical properties of molecular-level processes serve as a source of incipient pattern. By analogy, the evolution of consciousness can be thought of as depending in part on a competition between alternative variants in the microstructure of synaptic networks and/or the activity patterns they generate, some of which then serve as neural correlates of consciousness (NCCs).

Amphioxus neurocircuits, enhanced arousal, and the origin of vertebrate consciousness.

Gene expression studies have recently identified the amphioxus homolog of a domain comprising the combined caudal diencephalon plus midbrain, regions implicated in locomotory control and some forms of primary consciousness in vertebrates. The results of EM-level reconstructions of the larval brain of amphioxus, reviewed here, highlight the importance of inputs to this region for light and physical contact, both of which impinge on the same synaptic zone. The neural circuitry provides a starting point for understanding the organization and evolution of locomotory control and arousal in vertebrates, and implies that one of the tasks of midbrain-based consciousness, as it first emerged in vertebrates, would have been to distinguish between light and physical contact, probably sharp pain in the latter case, by assigning different qualia to each.

Amphioxus, motion detection, and the evolutionary origin of the vertebrate retinotectal map.

The axonal projection from the retina to the optic tectum maps visual information isomorphically from one to the other and serves as a model for the development of sensory maps more generally in the vertebrate brain. How or why this connection evolved is not known, nor why the midbrain is so important to the processing of visual information. Amphioxus is potentially informative here because its eye homolog, the frontal eye, also has a neural connection to a region of the brain now known to be homologous with the caudal diencephalon and midbrain.

The origin of dopaminergic systems in chordate brains: insights from amphioxus.

The basic anatomy of the central nervous system (CNS) is well conserved within the vertebrates and differs in significant ways from that of non-vertebrate chordates. Of the latter, amphioxus is of special interest, being the best available stand-in for the basal chordate condition. Immunohistochemical and gene expression studies on the developing CNS of amphioxus embryos and larvae are now sufficiently advanced that we can begin to assign specific neurotransmitter phenotypes to neurons identified by transmission electron microscopy (TEM), and then compare the distribution of cell types to that in vertebrate brains.

Interpreting amphioxus, and thoughts on ancestral chordate mouths and brains.

Amphioxus is increasingly important as a model for ancestral chordates. Nevertheless, it is secondarily modified in various ways, especially in the larva, whose small size has resulted in a rescaling and repositioning of structures. This is especially pronounced in the head region, where the mouth opens asymmetrically on the left side, leading to speculation that the mouth is secondarily derived, e.

The HMGA gene family in chordates: evolutionary perspectives from amphioxus.

High mobility group A proteins of vertebrates, HMGA1 and 2, are chromatin architectural factors involved in development, cell differentiation, and neoplastic transformation. Here, we characterize an amphioxus HMGA gene ortholog and analyze its expression. As a basal chordate, amphioxus is well placed to provide insights into the evolution of the HMGA gene family, particularly in the transition from invertebrates to vertebrates.

Locomotory control in amphioxus larvae: new insights from neurotransmitter data.

Amphioxus larvae have a midbrain-level locomotory control center whose overall organization is known from serial TEM reconstructions. How it functions has been a puzzle, owing to uncertainty as to the transmitters used by each class of neurons, but this has recently become clearer. We summarize what is now known, and correct past misconceptions: The large paired neurons at the core of the control center are glutamatergic, and hence excitatory, the commissural neurons are GABAergic, hence probably inhibitory, and both motoneurons and ipsilateral projection neurons are cholinergic, suggesting that the latter, a class of interneurons, may be derived evolutionarily from the former.

Echinoderm conundrums: Hox genes, heterochrony, and an excess of mouths.

Two issues relating to the translocation of anterior Hox genes in echinoderms to the 5' end of the Hox cluster are discussed: i) that developmental changes associated with fixation to the substratum have led to an acceleration of mesodermal development relative to that of ectoderm, resulting in a mismatch of anteroposterior registry between the two tissues and a larger role for mesoderm in patterning control, and ii) whether this helps explain the ability of some echinoderms to form separate mouths at different locations, one for the larva and one for the juvenile rudiment. Freeing the mesoderm from ectodermal influences may have encouraged morphogenetic innovation, paralleling the situation in tunicates, where an early genomic (or genomic and developmental) change has allowed the body to evolve in novel ways.

The Middle Cambrian fossil Pikaia and the evolution of chordate swimming.

Conway Morris and Caron (2012) have recently published an account of virtually all the available information on Pikaia gracilens, a well-known Cambrian fossil and supposed basal chordate, and propose on this basis some new ideas about Pikaia's anatomy and evolutionary significance. Chief among its chordate-like features are the putative myomeres, a regular series of vertical bands that extends the length of the body. These differ from the myomeres of living chordates in that boundaries between them (the myosepta) are gently curved, with minimal overlap, whereas amphioxus and vertebrates have strongly overlapping V- and W-shaped myomeres.

Development of the nervous system in hatchlings of Spadella cephaloptera (Chaetognatha), and implications for nervous system evolution in Bilateria.

Chaetognaths (arrow worms) play an important role as predators in planktonic food webs. Their phylogenetic position is unresolved, and among the numerous hypotheses, affinities to both protostomes and deuterostomes have been suggested. Many aspects of their life history, including ontogenesis, are poorly understood and, though some aspects of their embryonic and postembryonic development have been described, knowledge of early neural development is still limited.

Serial EM analysis of a copepod larval nervous system: Naupliar eye, optic circuitry, and prospects for full CNS reconstruction.

The medial eye and optic center of the first nauplius of Dactylopusia (=Dactylopodia) tisboides, a harpacticoid copepod, were reconstructed from serial EM micrographs. Axons from the eye project to a set of matching cartridges defined by glial cells processes, and input is then processed in sequence through two synaptic fields. A single class of local relay neurons provides the main pathway between these, subject to modulatory input from a class of densely stained neurons with distinctive dense terminals.

Head organization and the head/trunk relationship in protochordates: problems and prospects.

The fossil record has been an invaluable aid for reconstructing the major events of vertebrate evolution. There is no comparable record for protochordates, however, which severely limits our knowledge of their ancestral morphology, habits, and mode of life. The alternative is inference based on an interpretation of living protochordates but this is fraught with problems, not least being our own biases of what we think an ancestral chordate ought to look like.

The cholinergic gene locus in amphioxus: molecular characterization and developmental expression patterns.

The cholinergic gene locus (CGL), consisting of the vesicular acetylcholine transporter (VAChT)/choline acetyltransferase (ChAT) gene, encodes two specific cholinergic neuronal markers used extensively to study cholinergic transmission. In the present work, we isolated the amphioxus homologs of VAChT and ChAT and examined their expression during development. Analysis of the 5' untranslated region of VAChT and ChAT suggests that the splicing of the VAChT/ChAT mRNA has been evolutionarily conserved in amphioxus and mammals.

Basic features of the ancestral chordate brain: a protochordate perspective.

Basic features of the anterior nerve cord in amphioxus larvae are summarized to highlight its essential similarity with the vertebrate brain. Except for a pineal homolog, the amphioxus brain consists of a much simplified version of the ventral brainstem, including a region probably homologous with the hypothalamus, and a locomotory control center roughly comparable to the vertebrate tegmentum and reticulospinal system. Amphioxus has direct pathways for activating its locomotory circuits in response to mechanical stimuli via epithelial sensory cells, but this response is evidently modulated by inputs from diverse sensory-type cells located in the putative hypothalamic homolog, and from the lamellar body, the pineal homolog.

Prospective protochordate homologs of vertebrate midbrain and MHB, with some thoughts on MHB origins.

The MHB (midbrain-hindbrain boundary) is a key organizing center in the vertebrate brain characterized by highly conserved patterns of gene expression. The evidence for an MHB homolog in protochordates is equivocal, the "neck" region immediately caudal to the sensory vesicle in ascidian larvae being the best accepted candidate. It is argued here that similarities in expression patterns between the MHB and the ascidian neck region are more likely due to the latter being the principal source of neurons in the adult brain, and hence where all the genes involved in patterning the latter will necessarily be expressed.