An account is presented of the brain (paired cerebral, pleural, and pedal ganglia) of the nudibranch mollusc Tritonia diomedia. The major efferent nerve fibers are related to their nerve cell bodies and their functional roles identified as far as possible. The channels of sensory input relating to some of these neurons are given so as to provide an overall view of the organization of the brain.A standardized system of abbreviation and notation for the central ganglia, nerve trunks, and giant somata is propcsed. The system of reference is intended to provide a guide to the location in the ganglia of many of the smaller neurons of which the functional attributes are known, but which cannot be consistently recognized on visual criteria alone. A system of rectangular coordinates is proposed which is applied to the natural outline of the ganglia. I n addition, a system of cell notation is described which is independent of the co-ordinates used to define the position of the cell on the grid. Cells which by reason of their size, pigmentation, characteristic location and physiological attributes are consistently recognizable from animal to animal are numbered. Two principles were followed in numbering cells; (i) the series begins at unity in each ganglion; (ii) cell homologues in opposite ganglia are given the same number, but distinguished by prefixing the abbreviation for the ganglion in which they occur. It is considered that the system will facilitate the exchange of information between workers on the same species, and also benefit the comparison of neural organization of behavior in closely related forms. The brain is organized in an almost exactly bilaterally symmetrical manner. There are a few bilateral neural pathways, but the major functional routes are ipsilateral. A few motorneurons, which are uniquely identifiable anatomically, cause unique, discrete movements. Others are in small groups sharing overlapping or similar functions.
SUMMARYWe have investigated the roles played by numerous identified brain cells in initiating and controlling the coordinated sequence of movements of an instinctive escape-swimming sequence in an intact animal preparation of the nudibranch mollusc Tritonia diomedia. Intracellular electrical activity in different neurons has been correlated with the various phases of the behavior. We recognized four major stages in the response: (1) reflex local withdrawal; (2) preparation for swimming; ( 3 ) swimming; and (4) termination. We have located and studied brain cells whose activity is associated with the following aspects of swimming: withdrawal; elongation; triggering behavior; dorsal flexion; ventral flexion; and neurons which excite both dorsal and ventral flexor neurons simultaneously.We find that specific neurons play clearly defined and invariant roles in control of escape-swimming and that the neuronal circuitry underlying the coordination of the sequence is the same in different individuals of the species. Details of the neuronal circuitry and a number of the general functional attributes of interacting cell groups have been determined directly or inferred froin observations of cell to cell interactions. A preliminary model of the rieuronal apparatus which controls this behavior is discussed. The principal findings are: (1) a discrete group of electrically coupled neurons determines, by its output, whether or not escapeswimming will be executed; (2) the neuronal elements responsible for execution of the swimming stages of the sequence are maintained in an excited state for the required period, in part by a regenerative feedback system; ( 3 ) alternating bursts of impulses in functional antagonists are co-ordinated in part by reciprocal inhibition between them; and (4) termination of the sequence occurs abruptly a t a particular phase in the swimming cycle and appears to be an active neural process, rather than a simple running-down.
The germ cells of Polydora ciliata arise from cells lying just inside the ventral epithelium and later migrate into the gonads. They are released into the coelom when they are 30 μ in diameter and are transported posteriorly to the region of maturation. The eggs and sperm are shed through the modified nephridia of the region of maturation. The eggs are laid into a string of capsules apparently secreted by the nephridia and attached to the wall of the tube. The larvae take about a week to hatch and are released into the plankton at the three chaetiger stage. They spend some 6 weeks in the plankton before metamorphosis, and have grown to 17 or 18 segments. Monthly samples of the worms living in the London clay at Whitstable in 1956 showed that the population was bimodal with respect to size and the two groups of adults gave rise to two larval settlements in late March and May. The two groups of juvenile worms that developed from these larvae retained their identities through the summer and the following winter. There is no apparent morphological character by which the adults of each type may be recognized, the time of spawning and the lower fecundity of the early breeding worms being the only distinguishing features. It is suggested that these groups are the consequence of an overlap of two populations of P. ciliata from different centres of distribution, each being adapted in its breeding requirements to a different part of the temperature range.
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