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Feeding is the means by which an organism acquires the materials for building, maintaining and moving the vehicle that carries the next generation. Since nutrition is the main requirement of all living systems, feeding preceded by food-seeking behaviour, is a necessity of life. Organisms must feed to live and they also must work to feed. Evolution has played a role in influencing the eating and drinking behaviour of all species. It has selected for the mechanisms that motivate the organism's ingestion of nutrients and its selection. There are two fundamental aspects of feeding: energy balance and diet selection. Energy balance deals with how much animals eat in relation to their energy expenditures, whilst diet selection deals with mechanisms that allow omnivores to choose the appropriate nutrients. This chapter will deal with the former and Chapter 5 will be concerned with the latter issue.
Organisms regulate their nutritional intake according to shortand long-term energy needs. The mechanism responsible for this regulation, which involves the maintenance of a constant, optimal, internal environment, is called homeostasis. This concept was introduced by Claude Bernard (1879) in his discussion of ‘le milieu intérieur’ and the necessity of the organism keeping its internal environment at a constant, optimal level. (For a detailed discussion of the key components of homeostatic mechanisms and their character of operation, see Schulze (1995).) An example that illustrates the concept of homeostasis is that of the thermostat. After it is set for a certain temperature, it reacts to deviations from that temperature (the set point) by changing the environment so that such deviations are eliminated.
Motivational influences that are specifically social constitute the subject of this chapter. Social motives involve activities that affect interactions among organisms of the same species as well as organisms of other species. I will deal with only two issues in this short chapter: the formation and maintenance of social bonds between individuals; and that of pro-social and altruistic acts. The functional and proximate causal aspects of these phenomena will be considered in the analysis. The first and foremost social motive concerns the formation of the bond between an infant and its primary caretaker, usually the mother. The material in Chapter 3 deals with this issue from the perspective of the mother, whilst the material in this chapter complements it through an analysis of processes in the infant. The attachment of a human infant to its caretaker has biological roots. Both babies and adults are programmed by evolution to become attached in certain ways because the former is dependent upon the latter for survival. Attachment behaviours refer to a broad classification of behaviours that keep the infant in close proximity to an attachment figure. These behaviours include crying, clinging and approaching, as well as others produced when the infant is separated from the attachment figure.
Mammals live in a diverse array of habitats and social structures. The basic unit of the family is the mother and infant. However, our examination of material in Chapter 3 suggests that other conspecifics may be involved to some exent in infant caregiving. These caregivers can include the father, siblings or peers, or combinations of these conspecifics.
Drinking is the means by which an organism acquires fluids necessary for the normal functioning of the cells in the body. Water is the medium through which the chemical processes of the body operates. It is the largest component of the body and its volume must be defended within narrow limits. ‘The proportion of water to lean body mass (the body without fat) is essentially constant at 70%’ (Rolls & Rolls, 1982). The energy processes of the cell occur within a fluid medium. In Chapter 4, it was noted that feeding preceded by food-seeking behaviour, is a necessity of life. Similarly, drinking preceded by water-seeking behaviour is also a necessity of life. Living organisms are endowed with mechanisms that have been selected for, and which cause them to be motivated to seek and ingest water when their internal environment's water balance is disturbed. The animal's nervous system is supplied with information from a sample of body fluids, and drinking decisions are based upon the state of this sample. Just as the thermostat samples temperature at a site in a room, it is assumed that the drinking mechanisms sample the fluid environment at one or more sites.
Fluid regulation in living organisms represents a balance between intake and excretion of water. Each side of the equation consists of a ‘regulated’ and an ‘unregulated’ component. The regulated component represents factors which act specifically to maintain body fluid homeostasis (water balance). The primary factors that regulate water balance are thirst and pituitary secretion of the anti-diuretic hormone (ADH), which is also known as vasopressin (Verbalis, 1990).
When animals are exposed to aversive stimuli, particularly those of pain, they are likely to respond by either withdrawing from or attacking the source of the stimuli. ‘Pain is an anatomically developed sensory system genetically differentiated for survival and the defence of the body. Responses to painful stimuli either involve the skeletal musculature or are internal but they are experimentally quantified as escape and avoidance’ (Le Magnen, 1998, p. 4). Both types of responses to this source serve adaptive functions. In many species, specific escape mechanisms have evolved for dealing with physical danger. One of the simplest is the withdrawal reflex that removes the organism from damaging stimuli. When specific taste receptors are in contact with bitter substances, they result in a spitting reflex that protects the organism from ingesting possibly toxic substances that are generally associated with the bitter taste. In Chapter 5 the mechanisms by which rats learn to avoid smells and tastes that have previously been followed by illness were examined. Animals will also react to aversive stimuli with attack behaviour, particularly when escape is difficult. Such a reaction is particularly evident in feral animals.
TWO-FACTOR THEORY OF AVOIDANCE BEHAVIOUR
The standard situation or apparatus used to study responses to aversive stimuli is one in which a rat is placed in a shuttle box – a long narrow box divided in half by a partition. The floor of the box is a grid of steel rods that can can deliver a painful shock when activated by electricity. The rules of the experiment are as follows. The rat has a few seconds to cross the barrier over to the other side of the box.
This paper reports, for the first time, the existence of external digestion of decapod larvae by the common octopus, Octopus vulgaris (Mollusca: Cephalopoda), paralarvae. Zoeae of three crab species were externally digested, leaving a whole and empty exoskeleton. The attack sequence on these prey is also described, and divided into the same three phases (attention, positioning and seizure) already known for Sepia hatchlings.
Loxosomella tonsoria (Entoprocta: Loxosomatidae) was found associated with the polynoid polychaete Lepidonotus clava collected among the algae and debris scraped from the intertidal level at the rocky shore of Limens (Pontevedra, Spain). New features are added to the original description, together with the first description of its buds. Host specificity of the genus and distribution of this species are discussed.
Biomass measurements (AFDW) of both subtidal and intertidal macrofauna of the ‘Abra de Bilbao’ embayment were used within a pilot study in a four step strategy of analysis, which consisted of: (1) representing communities; (2) discriminating sites or conditions; (3) determining levels of ‘stress’; and (4) linking to environmental variables. Univariate (number of species, diversity indices) as well as multivariate (clustering, MDS, SIMPER, BIO-ENV) techniques from the PRIMER package were employed. Several data aggregation levels were proposed in the analyses with the aim of selecting cost-effective procedures. The results suggested a good integration of environmental conditions by subtidal samples, which were close to constant on all different taxonomic aggregation levels. The abiotic parameter which best ‘explained’ the biotic pattern of subtidal samples in the investigated area was turbidity measured as total suspension solids and nephelometric units on the water surface. In conclusion, we recommend monitoring rocky substrates by using more extensive biomass sampling surveys of subtidal areas followed by a less time-consuming treatment of the samples (identification by high taxonomic categories or by trophic groups).
Early life growth of the short-finned squid, Illex illecebrosus (Mollusca: Cephalopoda), was reconstructed from periodic increments deposited on the gladius structure of squid collected in the Gulf Stream/Slope Water front and the Scotian Shelf. The general effects of environmental changes and lifestyle transitions in the variation of size-specific gladius growth was investigated. The offshore growth was exponential and at least six size-specific growth phases were distinguished. Paralarval growth ended at approximately 10 mm gladius length (GL). An important change in growth at 30–40 mm GL was associated with the shift from the macroplanktonic to micronektonic habitats and the transition from the Gulf Stream to Slope Water. The approximation to the Scotian Shelf/Slope Water Front was suggested as related to a growth transition at 68 mm GL. A change to linear growth occurred after the nektonic lifestyle was attained on shelf waters (90 mm GL).
The occurrence of developmental abnormalities in two ovoviviparous littorinids (Littorina neglecta and a barnacle-dwelling form of L. saxatilis) inhabiting the barnacle zone at Peak Steel, Ravenscar on the north-east coast of England was investigated. In general developmental abnormalities appeared to be a rare occurrence in both taxa. However, L. neglecta was found to have a significantly greater proportion of deformed embryos in their brood pouches than L. saxatilis B (Figure 1). A distinct seasonal variation in the number of developmental abnormalities was observed in L. neglecta, and it is suggested this is a result of broods developing outside of the main reproductive season being more prone to deformity.
This communication reports on experiments, which studied the variation in competitive performance of juvenile cod (Gadus morhua L.) and their growth rates. The fish were held in groups of five in either summer or winter conditions and tested for their individual response to prey offered sequentially. There was marked individual variability. Fish that took the highest share of prey tended also to be those that took prey earlier than others. In winter conditions these fish were the largest, but in summer conditions size had no effect. There was a positive influence of the indices of competitive performance on individual growth rate but the relationship was not significant
Seasonal and spatial variations in the distribution and abundance of Ophiothrix fragilis larvae (2-arm to 8-arm stages) were determined from a series of plankton net tows in the Dover Strait in 1995 and 1996. Ophioplutei appeared in the plankton between June and September. Larval population was constituted by ophioplutei from different spatio–temporal origins. A major fraction of the ophioplutei observed in the Dover Strait might come from a population localized southward.
To examine the effects of submergence time, free-space availability, and larval supply on height-related differences in settlement density of Semibalanus cariosus, we monitored the daily settlement density in plots where all benthos were removed daily and in natural-community plots, which were left undisturbed, at two sites with different tidal heights: the centre and the upper limit of S. cariosus zone, during one settlement season. Settlement density was higher at the low-tidal site, where settler mortality was low, than at the mid-tidal site, where settler mortality was high. The between-site differences in settlement was determined by larval preference for height and not immersion time. The settlement preference for height was masked on days with high larval supply, since suitable settlement sites in lower height were saturated by settlers.
The caridean shrimp Rimicaris exoculata, Chorocaris chacei and Mirocaris fortunata, together with bathymodiolid mussels, dominate the vent fauna along the Mid-Atlantic Ridge. Vent shrimp show the characteristic reproductive patterns of caridean decapods. The gonads are paired organs overlying the digestive gland under the carapace. In the ovaries, the oogonia (∼20-30 μm diameter) proliferate in the germinal epithelium at the periphery of the gonad, developing into previtellogenic oocytes. The previtellogenic oocytes grow to 70-100 μm before undergoing vitellogenesis. The maximum size for mature oocytes ranged between 200 and 500 μm depending on the species and the sample. The oocyte size–frequency data show no evidence of synchrony in oogenesis at population level for any of the species studied. Mirocaris fortunata is the only species where gravid females are commonly collected. The brood is carried on the pleopods, and the number of eggs per female ranges from 25 to 503, with a mean egg length of 0.79±0.14 mm. There is a positive correlation between fecundity and body size, characteristic of crustaceans. One ovigerous C. chacei and two R.exoculata have been studied. The former was carrying 2510 eggs and the later 988 small eggs in an early stage of development. The fecundity of M. fortunata, C. chacei and R. exoculata is significantly higher than that of species from the Acanthephyra group collected in the north-east Atlantic.
Protozoan communities at three beaches on the Cantabrian Sea were studied. Four stations along a transect of 150 m at each beach were chosen for this study. The following parameters were measured: temperature, pH, oxidation–reduction potential, dissolved oxygen, conductivity, salinity, nitrate, phosphate, granulometry and amount of organic matter. Four groups of protozoa (ciliates, non-dinoflagellate flagellates, dinoflagellates and sarcodines) were analysed for their abundance and diversity, as well as the influence of the measured parameters on the communities. Temperature, pH, and dissolved oxygen were the main factors responsible for seasonal differences. Temperature increased at the beginning of spring and summer, coinciding with growth of the protozoan community. Flagellates and dinoflagellates increased during spring and early summer. The amount of organic matter subsequently rose. The final step of this succession was the development of the ciliate community. A comparison between the same areas in 1989 and in the present study was carried out.
A description is given of Alcyonidium reticulum sp. nov. (Bryozoa: Ctenostomatida), a smooth-surfaced species found encrusting intertidal stones and on Fucus serratus in south-west Britain. It is distinguished from two superficially similar congeners, A. gelatinosum and A. mytili.
Specimens of Opisthoteuthis grimaldii were video recorded in a shipboard aquarium and their behavioural traits, such as bottom resting, flat-spreading, web-inversion and ballooning response, were described. Ballooning, as defence response, has been recorded only for cirrate octopod families with an intermediate web (Cirroteuthidae: Stauroteuthidae), and this structure has been supposed to be essential for developing this behaviour. Members of the Opisthoteuthidae family lack an intermediate web, indicating that this is a common defence response in cirrates, expressed in a basic form in the Opisthoteuthidae, offering an illustration of how this behaviour can evolve.
Investigations were made on spatial and temporal variation, population structure, growth, recruitment and mortality in the intertidal flats of Kunsan (Korea, Yellow Sea) of an inarticulate brachiopod Lingula unguis. Statistical analysis indicates significant spatial differences and a significant relationship between density and ratio of sands to silts, suggesting that the distribution and density of this species were related to particle size. The population consisted of three year-classes, with a similar size composition each year, of which the second year-class was a major component. From estimation of parameters of growth monthly length–frequency was analysed by ELEFAN. Parameters of growth were estimated, using the modified von Bertalanffy growth function (VBGF) model incorporating seasonal variation in growth. The growth parameters are: L∞=44.00 mm SL, K=0.88 y−1, C=0.90, WP=0.48. The recruitment pattern shows two major recruitments per year. The maximum life span is estimated as 3.45 y. Total mortality (Z) by length-converted catch curve was estimated at 3.65 y−1. The population mainly occurred in fine sands and exhibited patchy distribution.
Larval development of Capitulum mitella (Cirripedia: Pedunculata) comprises six nauplius stages and a cyprid. Unilobed labrum, a larval trait of pedunculate and chthamalid barnacles, bears two groups of slender hairs and two pairs of strong teeth on the distal labral margin, and a pair of teeth on the lateral labral margin. Frontolateral horns extended under the anterior cephalic shield margin are diagnostic features through all nauplius stages. The posterior border of the cephalic shield bears a pair of distal cephalic shield spines in nauplius stages II and III, and a pair of long posterior shield spines in nauplius stages IV, V and VI. A hispid seta is consistently found at the fourth group of the antennal endopodite through stages II–VI. The dorsal thoracic spine, abdominal process and the paired posterior shield spines have numerous small spines. Morphological features such as the cephalic shield, labrum, abdominal process, antennules, antennae and mandibles in all nauplius and cyprid stages are illustrated and described. In this species, the numerical setations of the antennule are found to be beneficial for intra-specific identification of barnacle nauplius stages without the need for dissection.
In work performed during the period February–October 1996, about 47 species of ciliates were detected in the communities of Aufwuchs, or periphyton, colonizing glass substrates submerged in three sites in the Lagoon of Venice, two in the urban area of Chioggia and one at a mussel-farming site. The ciliate communities were composed of sessile forms of Peritrichia and Suctoria and of motile ciliates, belonging especially to Hypotrichia and Karyorelictea. Four species of the peritrich Zoothamnium and the suctorians Acineta tuberosa and Ephelota gemmipara are reported here for the first time in the Lagoon of Venice. The richest ciliate communities were recovered at 60 cm from the bottom. Processed by multivariate analysis, data of samplings performed up to day ten from the positioning of glass slides indicated significant differences in biotic and physico-chemical parameters, according to season. The first data did not give definite indications on the possibility of using ciliates colonizing solid surfaces to assess the quality of this peculiar environment.