The final effort of the CLIMAP project was a study of the last interglaciation, a time of minimum ice volume some 122,000 yr ago coincident with the Substage 5e oxygen isotopic minimum. Based on detailed oxygen isotope analyses and biotic census counts in 52 cores across the world ocean, last interglacial sea-surface temperatures (SST) were compared with those today. There are small SST departures in the mid-latitude North Atlantic (warmer) and the Gulf of Mexico (cooler). The eastern boundary currents of the South Atlantic and Pacific oceans are marked by large SST anomalies in individual cores, but their interpretations are precluded by no-analog problems and by discordancies among estimates from different biotic groups. In general, the last interglacial ocean was not significantly different from the modern ocean. The relative sequencing of ice decay versus oceanic warming on the Stage 6/5 oxygen isotopic transition and of ice growth versus oceanic cooling on the Stage 5e/5d transition was also studied. In most of the Southern Hemisphere, the oceanic response marked by the biotic census counts preceded (led) the global ice-volume response marked by the oxygen-isotope signal by several thousand years. The reverse pattern is evident in the North Atlantic Ocean and the Gulf of Mexico, where the oceanic response lagged that of global ice volume by several thousand years. As a result, the very warm temperatures associated with the last interglaciation were regionally diachronous by several thousand years. These regional lead-lag relationships agree with those observed on other transitions and in long-term phase relationships; they cannot be explained simply as artifacts of bioturbational translations of the original signals.

The cerebellar hypothesis of dyslexia posits that cerebellar deficits are associated with reading disabilities and may explain why some individuals with reading disabilities fail to respond to reading interventions. We tested these hypotheses in a sample of children who participated in a grade 1 reading intervention study (n = 174) and a group of typically achieving children (n = 62). At posttest, children were classified as adequately responding to the intervention (n = 82), inadequately responding with decoding and fluency deficits (n = 36), or inadequately responding with only fluency deficits (n = 56). Based on the Bead Threading and Postural Stability subtests from the Dyslexia Screening Test-Junior, we found little evidence that assessments of cerebellar functions were associated with academic performance or responder status. In addition, we did not find evidence supporting the hypothesis that cerebellar deficits are more prominent for poor readers with “specific” reading disabilities (i.e., with discrepancies relative to IQ) than for poor readers with reading scores consistent with IQ. In contrast, measures of phonological awareness, rapid naming, and vocabulary were strongly associated with responder status and academic outcomes. These results add to accumulating evidence that fails to associate cerebellar functions with reading difficulties. (JINS, 2010, 16, 526–536.)

The mechanical responses of several structures in the auditory and lateral-line systems of the sprat Sprattus sprattus(L.) and the herringClupea harengus L. to oscillatory pressure changes have been observed over a range of frequencies from 0·014 t0 2600 Hz. The pressures required to give constant responses of the bulla membrane, and the lateral recess membrane varied relatively little between 0·014 and 1000 Hz; above 1000 Hz the sensitivity fell quickly as frequency increased.

The orientations of the reflecting layers in the external surfaces of the herring have been found both by light-measurements on the fresh fish and by histological studies on preserved specimens.

The reflecting platelets which lie under the scales are orientated, with respect to the surface of the fish, in a similar way to the platelets found in the bleak and described earlier by Demon & Nicol (1965). However, on the curved dorsum of the fish, although the reflecting platelets are much more perpendicular to the sea surface than the scales on which they lie, these platelets are still inclined some 20° to the perpendicular. It is shown that, in this region, the fish reflects the fraction of the light striking the platelets which is sufficient to match the background against which the fish is seen. The platelets on the curved dorsum have the property of reflecting green light well if it falls obliquely on them but reflecting it poorly when it strikes them at angles close to normal incidence. On the broad flank of the herring the scales have reflecting platelets under most of their surfaces, and the individual scale has several distinctly coloured regions. When we look at any particular region of the flank of an intact fish we are always looking at several overlapping layers differing greatly from one another in their spectral reflecting properties.

It is the combination of the reflexions of several layers which gives the very bright silvery reflexions of the intact fish. A system of overlapping scales of this kind is needed even to reflect one waveband of light well over a wide range of angles of incidence.

The distribution of the visual pigments within the retina of the rudd, Scardinius erythrophthalmus, and dragonet, Callionymus lyra L., was studied by measuring the spectral absorbance of small areas of intact isolated retinas.

The rudd retina contains two visual pigments, VP 5O71 and VP5352. There is a progressive change in the relative proportion of these two pigments in the dorso-ventral plane, and there are no sharply defined zones each with a different pigment mixture. The proportion of the A1-based pigment (VP 5071) is greater in the dorsal part of the retina.

The maximal optical density of the pigment in the dorsal retina of the rudd was greater (about o·6) than it was in the ventral retina (about 0·3). The difference in the amount of light absorbed, however, is small compared to the intensity difference between the up-welling and downwelling light of the environment. When the relative proportion of the A1- and A2-based pigments was changed by altering the lighting conditions under which the animals were kept there was little or no change in the maximal optical density of the pigment.

In the dragonet the retina shows a clear division into a ventral pink zone (maximal optical density about 0·5) and a dorsal zone that is much paler (maximal optical density about 0·2). The relative spectral absorbance of the two regions is similar, although one is said to be a pure cone region, and the other to be dominated by rods.

The spectral absorbance curves for the intact retinas agreed well with results obtained with pigment extracts.

The newest chamber of a cuttlebone is always incomplete and full of a watery solution. This contains sodium and potassium in concentrations close to those of sea water.

The second newest chamber contains a gas space and sometimes contains liquid. This liquid is rarely localized at the siphuncular end of the chamber. It is not certain that liquid is initially extracted through the siphuncular wall of the chamber, although this is possible.

When liquid is pumped out of a newly formed chamber the salt which it initially contains is not left behind, either free or bound into the structure of the bone.

The 3rd to about the 10th newest chambers usually contain no visible liquid. But their siphuncular walls are permeable to liquid and liquid does enter these chambers when the cuttlebone becomes very dense.

At cuttlebone densities around 0·6, i.e. close to values which will make the cuttlefish neutrally buoyant, the older and more posterior chambers are almost full of liquid. This liquid can, however, be pumped out. When a cuttlebone has a density around 0·5 very little liquid can be seen in any of its chambers.

The pressure of gas within a newly ‘pumped out’ chamber is very low, but by the time a chamber has become the ninth newest the pressure of gas is close to the average value for the whole cuttlebone, i.e. about 0·8 atm.

The low pressure of gas found in the newest chambers is explained by the slowness with which gas diffuses into a space created by the active removal of liquid.

INTRODUCTION

The preceding paper (Allen, Blaxter & Denton, 1976) describes the development of the swimbladder-inner ear-lateral line system of the herring.

In the larval stage of herring the pro-otic bullae start to develop at a body length of about 18 mm. Between 18 and 30 mm the bullae become filled with gas. At 26 mm the lateral recess membrane starts to develop, becoming silvered at about 42 mm. The lateral line develops between 19 and 45 mm. The swimbladder silvers and contains gas from about 38 mm. The adult system is complete at 50–60 mm body length, some four months after hatching.

This paper concerns the system in its intermediate stages of development. It examines the problems of how the bullae are filled with gas and how the gas spaces are maintained. It gives evidence on the possible roles of the gas-filled spaces as hydrostatic pressure receptors and as aids to buoyancy and discusses the limitations of the system before it can gain or lose gas to a gas-filled swimbladder.

Direct observations on the reflecting layers in the tapetum of the spur dog Squalus acanthias confirm the orientations of the tapetal reflecting cells which were previously found histologically (Denton & Nicol, 1964). Surface views of the tapetum show the outlines of the exposed surface of the tapetal plates as ‘polygons’. The long axes of these polygons are parallel with the plane of the tissue on which they lie and the polygons are only inclined to this plane across their short axes. The angle to which they are inclined increases as we move towards the periphery of the eye.

Between April 1978 and July 1983 a total of 211 cuttlefish, Sepia officinalis L., was collected in the waters off Plymouth as part of a study of the cuttlebones (Denton & Gilpin-Brown, 1961). The following data were recorded for each specimen: sex, mantle length, the wet weights of the whole animal and the cuttlebone, and for females the weights of the eggs and nidamental glands. The presence of spermatophores in males was also noted. There are rather few records for the winter months; although Sepia were caught at this time they were often required for other studies. It is hoped to obtain more specimens for these months in later years.

INTRODUCTION

In earlier papers (Allen, Blaxter & Denton, 1976; Blaxter & Denton, 1976) an account was given of the development and structure of the swimbladder-bulla-lateral line system of the herring Clupea harengus L. and sprat Clupea sprattus (L.) and its function in the larval stage. In this paper we describe experiments on juveniles of these species in which the system is fully developed.

The behavioural experiments described in this paper were carried out to complement earlier physiological work on the acoustico-lateralis system of the clupeid fishes. Previous work, summarized by Blaxter, Denton & Gray (1981a), has shown that the sense organs of the clupeid utriculus are highly sensitive to alternating pressures. Since the utriculus signals the polarity and amplitude of each peak in a complex pressure wave form separately, independently of the rate of change of pressure over a range equivalent to a frequency band of 30–800 Hz, it is well suited to give information on brief noises. In the clupeids the lateral line is hydrodynamically connected with the ear and we have suggested that the signals arising from the inner ear and the neuromasts of the lateral line are used in conjunction to provide information about direction and range.

Herring larvae, when they are about 30 mm long, swallow air and fill their auditory bullae with gas. It has been shown earlier that this is associated with a great increase in sensitivity to changes in hydrostatic pressure. Here we show that it is also associated with the appearance of a startle response to noises.

At this stage of development, the utricular maculae are fully developed. However, the bullae are bounded by thin walls which are poorly calcified and the bulla membranes are thin and contain little elastic tissue. None the less the high-frequency cut-off in the mechanical responses to oscillating pressures is as high as that of the adult.

The reflecting layers in the external surfaces of salmon were studied during smoltification by making direct observations on the reflecting structures. The relationships between the reflecting layers in the epithelium external to the scales, in the tissues of the scales, and in the skin internal to the scales are given. Some fish, including fish blinded by pseudobranchectomy were reared under very abnormal lighting conditions and the processes of silvering and regeneration of scales followed on them. It was found that the pattern of silvering in smoltification and regeneration was not at all affected by lighting and that, although regenerated scales can easily be distinguished in other ways from those they replace, the orientations and reflectivities of their plate-lets are, with occasional exceptions, almost indistinguishable from those they replace.

INTRODUCTION

The herring Clupea harengus L. and sprat Sprattus sprattus (L.) are physostomatous teleosts with narrow ducts connecting the swimbladder to both the gut and cloaca. With other clupeoids these two species were of great interest to the anatomists of previous generations because of the further tubular connexions between the swimbladder and air-filled otic bullae close to the labyrinth of the inner ear. Together with the Ostariophysi, which have a chain of Weberian ossicles between the swimbladder and the inner ear, the clupeoids were thought to have enhanced hearing compared with many other teleosts as a result of coupling the ear to the swimbladder.

Despite such interest in the system the earlier literature is very fragmented, with the descriptions ranging over at least a dozen clupeoid species, and much of the work was done on fairly advanced juvenile or on adult fish. Ridewood (1891) examined the swimbladder-inner ear relationship in adult herring, pilchard Clupea pilchardus, sprat, shad C. alosa, twaite C. finta and anchovy Engraulis encrasicholus; Tracy (1920) made a similar study of the American Atlantic clupeoids – the shad Alosa sapidissima, alewife Pomolobus pseudoharengus, summer herring P. aestivalis, fall herring P. mediocris and menhaden Brevoortia tyrannus and O'Connell (1955) of the Pacific sardine Sardinops caerulea and anchovy Engraulis mordax. Wohlfahrt (1936) considered the total swimbladder-inner ear-lateral line relationship in 100–120 mm pilchards, recognizing the much less obvious connexion between the perilymph and the lateral line through a membrane in the skull. The presence of such a connexion had been suggested earlier by Tracy (1920).

The excess weight in sea water of the living tissues of Sepia officinalis (L.) is approximately balanced by the cuttlebone, which accounts for about 9·3% of the animal's volume. The density of cuttlebone varies around 0·6. The cuttlefish without its cuttlebone would be about 4% denser than sea water.

The chambers of the cuttlebone are independent of one another but liquids and gases are free to move within any one chamber.

Animals caught and studied fresh aboard ship exhibited a much less wide range of cuttlebone densities than those kept in an aquarium.

Specimens kept in aquaria vary greatly in buoyancy. These variations result from changes in density of the cuttlebone.

Cuttlebones differ not in the weight of dry matter per unit volume, which is always close to 38%, but in the amount of liquid they contain. A cuttlebone of density 0·7 contains about 30% liquid whereas a cuttlebone of density 0·5 contains about 10% liquid. The remainder of the cuttlebone contains gas, but this gas is at less than atmospheric pressure. The pressure of gas varies around 0·8 atmosphere. Within the duration of the experiments described here, the mass of gas per unit volume of bone remained almost constant whatever the bone's density. The pressure of gas is lower the less dense the cuttlebone. There can be no question of an evolution of gas expelling liquid from a bone when it becomes lighter. The constancy of the mass of gas within the cuttlebone is explained in terms of the slowness of diffusion of gases into the bone.

Squids (teuthoids) fall into two distinct groups according to their density in sea water. Squids of one group are considerably denser than sea water and must swim to stop sinking; squids in the other group are nearly neutrally buoyant. Analyses show that in almost all the neutrally buoyant squids large amounts of ammonium are present. This ammonium is not uniformly distributed throughout the body but is mostly confined to special tissues where its concentration can approach half molar. The locations of such tissues differ according to the species and developmental stage of the squid. It is clear that the ammonium-rich solution are almost isosmotic with sea water but of lower density and they are present in sufficient volume to provide the main buoyancy mechanism of these squids. A variety of evidence is given which suggests that squids in no less than 12 of the 26 families achieve near-neutral buoyancy in this way and that 14 families contain squids appreciably denser than sea water [at least one family contains both types of squid]. Some of the ammonium-rich squids are extremely abundant in the oceans.

Nautilus macromphalus Sowerby when freshly caught was close to neutral buoyancy having a weight in sea water of about 0–2% of its weight in air. The animals without their shells varied considerably in density but the volume of the shell was an approximately constant fraction of the total volume of the whole animal and whole animals were brought approximately to the same density by havingmore or less liquid inside the chambers of the shell. About 80 % of the gas space in the shell was used to support the weight of the shell itself in sea water.In an adult animal the centre of buoyancy was found to be about 6 mm above the centre of gravity, which made the animal very stable in its natural swimming position, a couple of about 350 g. cm being required to turn it through 90°. The pearly partsof the chamber walls were impermeable to sea water but the chalky and horny siphuncular tubes joining the septal necks were very porous. The most newly formed tenor so chambers were the only ones to contain liquids in appreciable volume and theydid this in diminishing amounts from the newest to the oldest. The watery liquids found within the chambers were always hypotonic to sea water and sometimes markedly so; they contained principally sodium and chloride ions. One animal was in the process of forming a new chamber, this incomplete chamber was completely full of liquidwith an osmolarity close to that of sea water but differing in composition from seawater.

Experiments are described which test the hypothesis that the cuttlefish controls the relative volumes of gas space and liquid within its cuttlebone by an osmotic mechanism acting across the siphuncular surface of the bone. When the animal is at the bottom of the sea it would maintain the gas space within the cuttlebone by balancing the hydrostatic pressure of the sea by an osmotic force between the liquid within the cuttlebone and the blood.

In cuttlefish kept for some weeks in shallow water in an aquarium all the liquid taken from the cuttlebone is almost isotonic with the animals’ blood.

In animals recently hauled from the bottom of the sea the cuttlebone liquid is markedly hypotonic to sea water and hence to the blood.

These lower osmotic concentrations are given chiefly by reduction in the concentrations of the sodium and chloride ions.

After bringing animals up from about 70 m to the surface of the sea the osmotic concentration of the cuttlebone liquid rises from about 75% of sea water some 40 min after starting to haul the trawl, to about 97% of sea water 6 h later. Extrapolation back to the time at which hauling began gives a concentration of salts close to that predicted by the osmotic hypothesis.

Whilst the cuttlebone liquid is increasing in ionic concentration the liquid deeper in the cuttlebone is hypotonic to that just inside the siphuncular surface. This is explained in terms of the slowness of exchange of salts along the narrow channels between the lamellae of the cuttlebone.