Cephalopod chromatophores: neurobiology and natural history

The chromatophores of cephalopods differ fundamentally from those of other animals: they are neuromuscular organs rather than cells and are not controlled hormonally. They constitute a unique motor system that operates upon the environment without applying any force to it. Each chromatophore organ comprises an elastic sacculus containing pigment, to which is attached a set of obliquely striated radial muscles, each with its nerves and glia. When excited the muscles contract, expanding the chromatophore; when they relax, energy stored in the elastic sacculus retracts it. The physiology and pharmacology of the chromatophore nerves and muscles of loliginid squids are discussed in detail. Attention is drawn to the multiple innervation of dorsal mantle chromatophores, of crucial importance in pattern generation. The size and density of the chromatophores varies according to habit and lifestyle. Differently coloured chromatophores are distributed precisely with respect to each other, and to reflecting structures beneath them. Some of the rules for establishing this exact arrangement have been elucidated by ontogenetic studies. The chromatophores are not innervated uniformly: specific nerve fibres innervate groups of chromatophores within the fixed, morphological array, producing 'physiological units' expressed as visible 'chromatomotor fields'. The chromatophores are controlled by a set of lobes in the brain organized hierarchically. At the highest level, the optic lobes, acting largely on visual information, select specific motor programmes (i.e. body patterns); at the lowest level, motoneurons in the chromatophore lobes execute the programmes, their activity or inactivity producing the patterning seen in the skin. In Octopus vulgaris there are over half a million neurons in the chromatophore lobes, and receptors for all the classical neurotransmitters are present, different transmitters being used to activate (or inhibit) the different colour classes of chromatophore motoneurons. A detailed understanding of the way in which the brain controls body patterning still eludes us: the entire system apparently operates without feedback, visual or proprioceptive. The gross appearance of a cephalopod is termed its body pattern. This comprises a number of components, made up of several units, which in turn contains many elements: the chromatophores themselves and also reflecting cells and skin muscles. Neural control of the chromatophores enables a cephalopod to change its appearance almost instantaneously, a key feature in some escape behaviours and during agonistic signalling. Equally important, it also enables them to generate the discrete patterns so essential for camouflage or for signalling. The primary function of the chromatophores is camouflage. They are used to match the brightness of the background and to produce components that help the animal achieve general resemblance to the substrate or break up the body's outline. Because the chromatophores are neurally controlled an individual can, at any moment, select and exhibit one particular body pattern out of many. Such rapid neural polymorphism ('polyphenism') may hinder search-image formation by predators. Another function of the chromatophores is communication. Intraspecific signalling is well documented in several inshore species, and interspecific signalling, using ancient, highly conserved patterns, is also widespread. Neurally controlled chromatophores lend themselves supremely well to communication, allowing rapid, finely graded and bilateral signalling.     

Cephalopod Cognition in an Evolutionary Context: Implications for Ethology

What is the distribution of cognitive ability within the animal kingdom? It would be egalitarian to assume that variation in intelligence is everywhere clinal, but examining trends among major phylogenetic groups, it becomes easy to distinguish high-performing ‘generalists’ – whose behavior exhibits domain-flexibility – from ‘specialists’ whose range of behavior is limited and ecologically specific. These generalists include mammals, birds, and, intriguingly, cephalopods. The apparent intelligence of coleoid cephalopods (squids, octopuses, and cuttlefish) is surprising – and philosophically relevant – because of our independent evolutionary lineages: the most recent common ancestor between vertebrates and cephalopods would have been a small wormlike organism, without any major organizational structure to its nervous system. By identifying the cognitive similarities between these organisms and vertebrates, we can begin to derive some general principles of intelligence as a biological phenomenon. Here, I discuss trends in cephalopod behavior and surrounding theory, and suggest their significance for our understanding of domain-general cognition and its evolution.     

Cubozoan jellyfish: an Evo/Devo model for eyes and other sensory systems

Cnidaria are the most basal phylum containing a well-developed visual system located on specialized sensory structures (rhopalia) with eyes and statocyts. We have been exploring the cubozoan jellyfish, Tripedalia cystophora. In addition to containing simple photoreceptive ocelli, each rhopalium in Tridedalia has a large and small complex, camera-type eye with a cellular lens containing three distinct families of crystallins which apparently serve non-lenticular functions. Thus, Tridpedalia recruited crystallins by a gene sharing strategy as have mollusks and vertebrates. Tripedalia has a single Pax gene, PaxB, which encodes a structural and functional Pax 2/5/8-like paired domain as well as an octapeptide and Pax6-like homeodomain. PaxB binds to and activates Tripedalia crystallin promoters (especially J3-crystallin) and the Drosophila rhodopsin rh6 gene in transfection tests and induces ectopic eyes in Drosophila. In situ hybridization showed that PaxB and crystallin genes are expressed in the lens, retina and statocysts. We suggest from these results that an ancestral PaxB gene was a primordial gene in eye evolution and that eyes and ears (mechanoreceptors) may have had a common evolutionary origin. Thus, the numerous structural and molecular features of Tridpalia rhopalia indicate that ancient cubozoan jellyfish are fascinating models for evo/devo insights into eyes and other sensory systems.     

How biologists conceptualize genes: an empirical study

Philosophers and historians of biology have argued that genes are conceptualized differently in different fields of biology and that these differences influence both the conduct of research and the interpretation of research by audiences outside the field in which the research was conducted. In this paper we report the results of a questionnaire study of how genes are conceptualized by biological scientists at the University of Sydney, Australia. The results provide tentative support for some hypotheses about conceptual differences between different fields of biological research.     

Writing and speaking: communication for biologists

A workshop for university biology teachers examined the teaching to undergraduates of writing and speaking. There was agreement that the communication of results and ideas was an integral part of scientific method. Proposals for the teaching of different aspects of communication during the undergraduate course are given and an Appendix lists some material which has been used with, and found suitable for biology students.     

Dilemmas in conservation for applied biologists

Prevention of the use of DDT has been made the target of a powerful propaganda drive in certain prosperous countries because, it is stated, DDT is a danger to man and harms wild life. On the other hand, DDT is by far the most economical, effective and safe insecticide for many uses, particularly for protecting men from certain insect-borne diseases and for enabling cotton to be grown in poor countries. Some risks can be reduced by eliminating those uses of DDT for which adequately safe, economical and effective substitutes exist, whether chemical or not; other risks can be reduced in other ways. The known risks to men are trivial, except when DDT concentrate is deliberately drunk, and the scare is made up of unknown risks -which could equally exist with any object or material, new or old. Risks to wild life have been greatly exaggerated and scares depending on falsehoods have become current. The postulated threat of progressive accumulation of DDT along a long food chain is not adequately supported by evidence, much of which has been misinterpreted. Thus the main dilemma is how to balance the great and undoubted benefits of DDT to millions of men, women and children against harm to wild life, sometimes genuine and remediable and sometimes dubious. People who campaign for banning have possibly failed to recognize this dilemma. On the other hand, they may have made a deliberate choice in favour of wild life. In that case, to be logical, they should also oppose all other means of preventing premature death of other people, which they might justify as a means of postponing over-population. The use or abuse of DDT is a minor component in the rise of the worl's population.