The relation between feeding and reproduction in Anasterias minuta (Asteroidea: Forcipulata)

Anasterias minuta is an oral-brooding sea star and one of the main predators of intertidal benthos in Patagonia, Argentina. The aims of this study were to: (1) assess the temporal pattern of feeding and brooding, (2) evaluate the effect of body size, tidal height and temperature on feeding and brooding, (3) evaluate temporary changes in digestive and sexual organs and (4) investigate the relationships between the predatory and brooding activities. Females do not feed during the brooding period. Spawning proceeded during March. The brooding period was from March to November, with a maximum in May. The population feeding rate varied seasonally, with a maximum in December and a minimum in May. The feeding rate was positively related to water temperature and sea star size; but not with tidal height. An annual cycle exists for pyloric caeca weight in both sexes; lowest values were found during winter, due to brooding female starvation, and a possible decrease in feeding rate of non-brooding females and males. No inverse relationship was found between the pyloric caeca and gonad weights in females, but a lag of 1 month was observed in males, being the maximal pyloric caeca development before the maximum gonad weight, suggesting nutrient translocation.     

The biochemical composition, energy content, and chemical antifeedant defenses of the common Antarctic Peninsular sea stars Granaster nutrix and Neosmilaster georgianus

The sea stars Granaster nutrix and Neosmilaster georgianus are conspicuous members of benthic communities along the Antarctic Peninsula. An analysis of the proximate composition of somatic body components of nonreproductive adults indicates the nutrient storage organs (pyloric caeca) are rich in both protein (60.7 and 60.6% mean dry wt, respectively) and lipid (25.4 and 29.8% mean dry wt, respectively). Body-wall tissues, while containing small inconspicuous skeletal ossicles, are also comprised of significant levels of organic matter (33.5 and 55.7% mean dry wt, respectively), attributable primarily to protein. Both the pyloric caeca and body-wall tissues are relatively rich in energy (mean energy levels=24.8 and 26.5 kJ g⁻¹ dry wt; 8.4 and 14.1 kJ g⁻¹ dry wt, respectively). Despite the availability of these nutrients and energy neither sea star is preyed upon by the sympatric omnivorous sea star Odontaster validus, a common predator of other Antarctic sea stars. Laboratory feeding bioassays indicate that O. validus rejects live intact individuals and body-wall tissues of both sea star species while readily consuming dried krill. Alginate food pellets containing a krill powder and tissue level concentrations of organic methanol extracts of body-wall tissues were also rejected by O. validus. Moreover, the copious mucus released from the body wall of N. georgianus was deterrent in O. validus food pellet bioassays. Thus, both sea stars evidently possess defensive secondary metabolites that defend against predation and are likely to play a role in mediating materials and energy transfer in the Antarctic benthos.     

The effect of feeding or starvation on resource allocation to body components during the reproductive cycle of the sea urchin Sphaerechinus granularis (Lamarck)

To determine the effects of feeding or starvation on resource allocation to body components during the reproductive cycle of Sphaerechinus granularis, sea urchins were placed in laboratory tanks and either fed ad libitum or starved during two different periods of their biological cycle, i.e. the mature stage and the recovery stage. The urchin growth was monitored over the whole experimental period, the gonad, gut, lantern indices and organic matter levels of different organs were determined at the end of the experiment. During the mature stage sea urchins in good nutritional conditions did not increase in size, but allotted energy to gonad production and stored reserves in body wall. Limiting food stopped the gonadal growth without complete regression. During the recovery period food allowed somatic growth, i.e. test growth and the storage of reserves in gonad somatic cells. This somatic production did not occur under food-limited conditions and the resources allotted for survival and maintenance were taken from different body components.     

Effect of the degree of autotomy on feeding, growth, and reproductive capacity in the multi-armed sea star Heliaster helianthus

The term autotomy refers to the process by which some species lose limbs or parts of limbs in response to adverse biotic or abiotic conditions, as for example, predation or abnormally high temperatures. The multi-armed sea star Heliaster helianthus is a key predator of the intertidal and the shallow rocky subtidal communities of north-central Chile. Natural populations of this sea star have been found with up to 60% of the individuals showing some degree of autotomy. The present study evaluated the effects of autotomy on feeding rate and growth of juvenile and adult H. helianthus after experimentally induced autotomy of 17% and 33% of their arms, as well as on the energy content of the pyloric caeca and gonads of adults during the reproductive period. Experimental juvenile sea stars were maintained and fed in the laboratory over a period of five months and adult sea stars for one month, Intact individuals were maintained as parallel controls. The results showed that juveniles undergoing 33% autotomy decreased their feeding rates, and as a consequence showed lowered net individual growth. In contrast, adults with 17% and 33% autotomy showed marked reductions in feeding. The results showed that autotomized adults had between five and seven times lower contents of carbohydrates, lipids, and proteins (and thus energetic content) in their pyloric caeca and gonads. The loss of the arms not only decreased the capacity for feeding in sea stars, but also allocated energy away from growth and reproduction into the process of regeneration of arms. This suggests that autotomy reduces the fitness of H. helianthus. Growth was reduced in the juveniles, while adults became limited in their ability to store energy which then limited their reproductive potential. Finally, based on the important effect of autotomy on reducing the feeding capacity of H. helianthus, the role of this sea star as a predator in the environment may be strongly affected.     

Rate of regeneration of two arms in the field and its effect on body components in Luidia clathrata (Echinodermata: Asteroidea)

The incidence of arm regeneration, effects of arm regeneration on non-regenerating body components and rate of arm regeneration were examined under field conditions. Approximately 60% of Luidia clathrata found in the upper portion of Tampa Bay, Florida were regenerating one or more arms. Individuals with two arms amputated at the disk edge took 380 days to regenerate completely. Both arms of an individual regenerated at the same rate. Length of regenerating arms initially increased faster than dry mass. No difference was found between non-regenerating and regenerating individuals for the dry mass of the body wall, pyloric caeca, and gonad of a non-regenerating arm. The absolute masses of the gonad and pyloric caeca were low through out the year possibly indicating a low food environment. Based on the rate of regeneration, arm loss just prior to the reproductive season may reduce the reproductive potential of this species.     

Autotomy reduces feeding, energy storage and growth of the sea star Stichaster striatus

We evaluated the effect of autotomy on feeding, energy storage and growth of juvenile Stichaster striatus kept in the laboratory for five months with a limited supply of the mussel Semimytilus algosus. Autotomy strongly decreased feeding, energy storage and growth. Intact juveniles showed a ∼ 3 fold higher feeding rate than autotomized individuals throughout the experiment. Intact juveniles also had a higher (∼ 5 fold) energy content per pyloric caeca in each arm. This was mainly due to higher lipid content, the main proximate constituent of pyloric caeca. Intact juveniles showed a greater growth rate and reached a greater size than autotomized individuals, more evident for underwater mass than radius length. The reduced capacity to feed reduced energy intake in autotomized individuals. However, low energy reserves along with low growth in autotomized sea stars, support the hypothesis that juveniles of this species allocate energy to regeneration to the detriment of growth. This was also supported by the ∼ 25% of arm length regeneration after 5 mo. Remaining small could increase risk of lethal predation, however, S. striatus may reduce predation risk by using crevices and kelp holdfasts as refuges from predators. Given the strong impact of autotomy on feeding, regeneration of arms to recover full capacity to forage and grow seems a better strategy for juvenile S. striatus, than merely growing.     

Asexual reproduction and somatic growth of the fissiparous sea star Allostichaster polyplax in New Zealand

Allostichaster polyplax is a fissiparous (capable of reproducing asexually by fission) sea star found in intertidal and shallow subtidal habitats in New Zealand and Australia. A localized population of A. polyplax occurred at a density of 2.7?individuals?m^?2 in an intertidal boulder field at Otago Harbour, South Island, New Zealand. The percentage of sea stars undergoing fission, recorded from January 2010 to March 2011, peaked in January (midsummer) in both years. The frequency of fission decreased for larger sea stars (mean arm length, R?>?25?mm), suggesting an ontogenic shift from asexual to sexual reproduction. Effects of fission (i.e. arm regeneration) and food availability (small gastropods) on growth, storage and sexual reproduction were examined in a factorial laboratory experiment. There was no effect of arm regeneration or food on total arm growth. Storage capacity, measured as pyloric caeca index (PCI), was greater in full-armed than regenerating sea stars, and greater in fed than starved individuals. Fed individuals had a similar PCI as field-collected ones. Gonads were observed in two individuals (males) in the full-armed and fed treatment. Fission did not occur during the experiment. Feeding rate on snails was greater for full-armed than regenerating sea stars. Water temperature had a weak positive effect on the feeding rate of regenerating sea stars. Our findings suggest that fission and subsequent growth of ramets can sustain demographically stable and geographically isolated populations of A. polyplax. The role of sexual reproduction in population dynamics and connectivity of this species (and fissiparous sea stars in general) remains equivocal and requires further research on genetic differentiation within and among populations.     

Physiological correlates of growth and condition in the yellow perch (Perca flavescens)

This study on yellow perch (Perca flavescens) examines a series of enzymatic markers and the relative weights of pyloric caeca and visceral lipids, their response to changes in feeding regime and their potential use to infer recent changes in growth rate and fish condition. Fish were exposed to four different feeding regimes for 12 weeks resulting in specific growth rates ranging from 0.3% to 3.5% (%/day). Growth and condition responded rapidly to changes in ration and the weight of pyloric caeca and visceral lipids reflected increased feed intake. Growth rate was correlated with muscle citrate synthase and caecal nucleoside-diphosphate kinase activities, whereas condition was correlated with muscle citrate synthase and lactate dehydrogenase activities and with caecal glucose-6-phosphate dehydrogenase activity. Results showed that enzyme activities and biometric parameters responded rapidly to increased feed intake, but the response was slower when food intake decreased. Plateaus were attained for both condition and visceral lipid index, but the relative weight of pyloric caeca continued to increase throughout the experimental period. Results from this study could, in principle, be used to infer recent growth and energy status in wild yellow perch and thus provide an indicator of food availability in their environment.     

The annual pyloric cycle of Asterias rubens L. (echinodermata: Asteroidea)

Observations of biochemical changes in the pyloric caeca of Asterias rubens L. in different nutritional states and at different stages of the reproductive cycle show the existence of an annual cycle in the pyloric caeca. This pyloric cycle is closely related to that of the gonads. During the gonadal rest (May to September, when the gonads are barely visible) the pyloric caeca grow and store glycogen. During gonadal activity (October to April), they progressively autolyse and first accumulate, then liberate, lipid reserves. This winter pyloric autolysis provides the gonads with the structural material necessary for their annual growth. The summer pyloric growth must be interpreted as the accumulation of the structural material. Under normal conditions, during gonadal rest, the pyloric glycogen reserves ensure the survival of the animal. Pyloric lipid reserves appear only at the beginning of gonadal activity. Their synthesis is intimately related to the start of the gonadal activity of which they are the principal energy source. We believe that the pyloric cycle is under control of hormones that could be transported through the haemal system.The pyloric caeca are the only reserve organs of starfish. Neither the other digestive organs nor the body wall accumulate energy reserves. During starvation of medium duration (2 to 4 weeks) the pyloric caeca suffice to ensure the survival of the animal by liberating their reserve material and simultaneously starting to autolyse. During prolonged starvation, general autolysis occurs and the size of the starfish diminishes.     

The feeding, growth, and energetics of two rocky intertidal predators (Pisaster ochraceus and Nucella canaliculata) under water temperatures simulating episodic upwelling

Although most upwelling regions are marked by strong fluctuations in water temperature, few studies have examined how episodic cold-water events affect the physiology and ecology of benthic marine invertebrates. I tested the hypothesis that upwelling-related variation in water temperature regulates the feeding, growth, and energetics of two rocky intertidal predators, the sea star Pisaster ochraceus (Brandt, 1835) and the whelk Nucella canaliculata (Duclos, 1832). Sea stars and whelks were maintained in laboratory tanks at a constant 9 °C, a constant 12 °C, and a treatment that simulated the Oregon coast upwelling regime by cycling between 14-day periods of 12 and 9 °C. Early in the experiments, sea stars and whelks held at 9 °C consumed about 30% fewer mussels (Mytilus trossulus) than those in warmer tanks. Despite lower consumption by whelks in colder tanks, 9 and 12 °C individuals attained the same final size. Similarly, sea stars in 9 °C tanks showed greater growth per gram of mussel tissue consumed than individuals held at 12 °C. These results suggest that reduced consumption under colder conditions was balanced by reduced metabolic costs. Moreover, there appeared to be an energetic advantage to living in the temperature regime characteristic of intermittent upwelling. Sea stars alternately exposed to 12 and 9 °C had a significantly higher growth rate, conversion efficiency, and storage of reserves in the pyloric caeca than individuals in the constant 12 °C tanks. Whelks maintained under fluctuating temperatures tended to grow faster than those held at constant 12 or 9 °C, although this trend was not statistically significant (p=0.069). These results suggest that benthic consumers experiencing cyclic temperatures may feed intensely during periods of warmer water while benefiting from reduced metabolic costs during cold-water intrusions. Because the fecundity of Pisaster and Nucella is a function of energy stored during the upwelling season, interannual variability in upwelling patterns could alter the reproductive output of these species.     

Phenotypic flexibility of digestive system in Atlantic cod (Gadus morhua)

This study examined the restoration of the digestive capacity of Atlantic cod (Gadus morhua Linnaeus) following a long period of food deprivation. Fifty cod (48 cm, 1 kg) were food-deprived for 68 days and then fed in excess with capelin (Mallotus villosus Müller) on alternate days. Ten fish were sampled after 0, 2, 6, 14 and 28 days and the mass of the pyloric caeca, intestine and carcass determined. Two metabolic enzymes (cytochrome c oxidase and citrate synthase) were assayed in white muscle, pyloric caeca and intestine, and trypsin activity was measured in the pyloric caeca. A delay of 14 days was required before body mass started to increase markedly, whereas most of the increase in mass of both the pyloric caeca and intestine relative to fish length occurred earlier in the experiment. By day 14, the activities of trypsin and citrate synthase in the pyloric caeca as well as citrate synthase in the intestine had reached maxima. The growth of the digestive tissues and restoration of their metabolic capacities thus occur early upon refeeding and are likely required for recovery growth to take place. The phenotypic flexibility of the cod digestive system is therefore remarkable: increases in trypsin activity and size of pyloric caeca resulted in a combined 29-fold increase in digestive capacity of the fish during the refeeding period. Our study suggests that Atlantic cod are able to cope with marked fluctuations in food availability in their environment by making a rapid adjustment of their digestive capacity as soon as food availability increases.     

Level,content, and caloric equivalents of the lipid,carbohydrate, and protein in the body components of Luidia clathrata (echinodermata: asteroidea: platyasterida) in Tampa bay

The body components of the primitive platyasterid starfish, Luidia clathrata (Say) have been characterized according to season, reproductive, and nutritional condition. The animal has been ‘reconstructed’ in terms of the wet and dry weights, the weight of organic matter, and the caloric equivalents of the body components in early summer (when there are no gonads and the pyloric caeca are small), in autumn (just prior to initiation of gonadal development and at maximal caeca size), and in early spring (at maximal gonad size). The body wall is always the most significant component in terms of wet and dry weights. In terms of organic matter and calories, the pyloric caeca are the most significant compartment in the fall and, together with the gonads, in the spring. On starvation for one month, the decrease in the size of the pyloric caeca is calculated to have produced 2.048 kcal. The energy requirement over the period, calculated from Q_o2 values, is 2.112 kcal. There was no change in biochemical composition of the pyloric caeca with starvation, suggesting that energy production came about through cell destruction rather than preferential utilization of cellular nutrient stores. Lack of extreme change in the composition of the pyloric caeca with size through the year also suggests that energy deposition in the organ is primarily by change in cell number and not by cell size.     

Reproductive patterns within sub-populations of Lytechinus variegatus (Lamarck) (Echinodermata: Echinoidea)

The annual reproductive cycle of the sea urchin Lytechinus variegatus (Lamarck) was examined in individuals collected from four habitats in the Anclote estuary near Tarpon Springs, Florida. Two periods of advanced gametogenic activity were apparent, but this bimodal pattern was not necessarily reflected by gonad growth curves. Differences in reproductive events among sub-populations were related primarily to patterns of gonad growth. Reproductive synchrony present during spring became less apparent as spawning and recovery within the sub-populations took place at differential rates during late-spring and summer. The final spawnout in late summer returned the sub-populations into gametogenic synchrony.     

The effects of various diets on chitinase and ß-glucosidase activities and the condition of cod, Gadus morhua (L.)

Groups of cod, Gadus morhua (L.), were fed exclusively on fish, crustaceans, or crustacean shells for a period of 3 weeks. Chitinase and ß-glucosidase activities were measured in enzyme extracts of stomach contents, stomach tissue, pyloric caeca, intestinal contents, and intestine tissue, and compared to the enzyme activities of control fish starved over the same period. Fulton's condition factor K , liver lipid content and liver water content were determined to estimate the effects of the diets on the condition of cod.
general, the highest chitinase activities were measured in samples of cod that had been fed on whole crustaceans. In this feeding group, there was also a remarkable increase of activity by a factor of four to eight in the pyloric caeca, compared to the group fed on fish and the control group, respectively.
Measurements of ß-glucosidase activity revealed no similar dependence on food quality, ß-glucosidase occurrence seemed to be mainly restricted to the pyloric caeca and the intestine of cod.     

Patterns of sperm-specific histone variation in sea stars and sea urchins: primary structural homologies in the N-terminal region of spermatogenic H1.

An electrophoretic characterization of histones from pyloric caeca, testes, and sperm of Asterias vulgaris revealed a sperm/testes-specific variant of histone H1 significantly larger than its somatic counterpart from pyloric caeca. Additional proteins were observed in H1 regions of acetic acid-urea polyacrylamide gels in testicular extracts. Sperm or testis-specific variants of H2B observed in sea urchins were not found in the sea star. Evidence presented suggests that sperm- or testes-specific H1 species of intermediate mobility may arise from a single, slow-migrating H1 species (SpH1). Although an increase in nonspecific DNA binding by nuclear proteins must occur during the process of spermatogenesis, different organisms exhibit various patterns of sperm-specific protein mediating differential binding during the process. Sperm-specific variants of both H1 and H2B histones are observed in sea urchins, while the only variant observed in sea stars during spermatogenesis is SpH1. Sequencing of the N-terminus of SpH1 from A. vulgaris revealed a repeating tetrapeptide in residues 3-6 and 8-11 (Ser-Pro-Arg-Lys and Ser-Pro-Lys-Lys, respectively), homologous to repeats in the N-termini of sperm-specific H1s from sea urchins. Primary structure within critical, variable regions of molecules responsible for nonspecific DNA binding appear conserved in many organisms. The occurrence of repeating tetrapeptides in SpH1 and other DNA binding proteins suggests that such domains may function similarly in various chromatins undergoing regulated or reversible condensation.     

Mechanisms of digestion in the marine herbivore, the luderick, Girella tricuspidata (Quoy and Gaimard)

The effect of feeding frequency on the mechanisms of digestion in the mucosa and contents of the gastrointestinal tract of the luderick, Girella tricuspidata (Quoy and Gaimard), was determined. Use was made of the variation in utilization frequency of Enteromorpha intestinalis by C. tricuspidata fed according to two different regimes to determine the importance of these digestive mechanisms in algal digestion.
The contents of the oesophagus and stomach of fish fed ad libitum were consistently acidic whilst those from fish deprived of food were highly variable and more alkaline. The contents of the pyloric caeca. intestine and rectum were slightly alkaline in both groups.
The number of microorganisms found in the digestive tract increased with a reduction in food availability. There was no significant cellulase activity in any of the animals tested.
Amylase and lipase were found in the pyloric caeca, intestine and rectum, but were not present in the oesophagus or stomach. Lipase activities were highly variable. Although some protease was found in the posterior portion of the gut, the majority of the activity occurred in the oesophagus and stomach.
Generally, amylase, lipase, and protease activities were unaffected by a change in the feeding regime of the fish from being fed ud libitum to being fed for a period of 24 h every 5 days. It is concluded that a reduced activity of enzymes that hydrolyse algal cell contents is not the reason for the lower utilization efficiency of food-deprived fish.
It is clear that the only difference in digestive mechanisms between fish fed ad libitum and fish fed infrequently is the variation in pH of the foregut. It is considered likely that the increased acidity of the foregut of animals fed ad Iibilum facilitates hydrolysis of the algal cell walls and allows greater utilization of algae by these fish.     

The accumulation,distribution, and toxicological effects of dietary arsenic exposure in lake whitefish (Coregonus clupeaformis) and lake trout (Salvelinus namaycush)

A 20-day experiment was conducted to compare the accumulation, distribution, and toxicological effects of dietary As, as arsenate, in lake whitefish (LWF, Coregonusclupeaformis) and lake trout (LT, Salvelinus namaycush). Results of this experiment were used to design an experiment of longer duration in which one of the fish species was selected and exposed three times per week to lower dietary As doses. In the present study each treatment group was exposed to a combination of one of three doses of As (0, 100, or 1000 μg As/g) and one of two types of diet, no brine shrimp (NS) or with brine shrimp (WS) for a total of eight dosing events. Brine shrimp were added to determine whether their presence enhanced consumption of As-contaminated food. Modified feeding behavior occurred in both fish species fed As contaminated diets, with the exception of the 100 μg As/g NS food. Brine shrimp addition did not affect feed consumption of the As contaminated diets. Significant As accumulation occurred in stomach, pyloric caeca, intestine, liver, kidney, and gallbladder, but not in bile or muscle. As exposure did not have a significant effect on hepatic and renal metallothionein concentrations. Concentrations of lipid peroxides were only significantly elevated in the plasma of LT fed the 1000 μg As/g WS food. Liver somatic indices decreased significantly in both species, whereas hematological parameters were not affected in either species. Histological lesions occurred in gallbladder, liver, kidney, pyloric caeca and intestine from LWF. These lesions were not observed in LT; however, gallbladders were not examined in this species. Weight gain was lower in both species fed As contaminated diets, however, condition factors were not affected.     

A non‐lethal method for estimation of gonad and pyloric caecum indices in sea stars

Abstract. Gonad and pyloric caecum indices are widely used indicators of reproductive effort and nutritional condition in asteroids. Current methods of quantification generally require sacrificing multiple animals and the resulting reduction in local sea star density could have an unintended impact on benthic communities. Using the intertidal sea star Pisaster ochraceus, we developed and tested a method for estimating organ indices through the non‐lethal sampling of single arms. Indices estimated via dissections of single arms accurately predicted the values obtained by sacrificing whole animals. In laboratory and field trials, we compared two methods of sampling single arms: (1) arm removal, and (2) organ extraction through an incision (without arm removal). Two years after these treatments, organs had regenerated in the affected arms of most sea stars, but were small relative to those in unmanipulated arms. The extent of organ recovery did not differ between the two treatments, but sea stars in the arm removal group were relocated in the field more frequently than those in the arm incision group. Our results suggest that sampling via the removal of single arms can be an effective, non‐lethal method for estimating gonad and pyloric caecum indices in asteroids.     

Androgen metabolism in somatic and germinal tissues of the sea star Asterias vulgaris.

1. Cell-free homogenates of male and female pyloric caeca, body wall, testis and ovary were incubated with radiolabeled 3H-androstenedione. 2. Pyloric caeca had highest rates of androstenedione conversion. The predominant metabolites in the pyloric caeca were testosterone, 5 alpha-androstane-3 beta, 17 beta-diol and 5 beta-androstane-3 beta, 17 beta-diol. 3. In body wall, testicular and ovarian homogenates, androstenedione was converted primarily to testosterone and also to 5 alpha-androstanedione and epiandrosterone. 4. Qualitative and quantitative differences in androgen metabolism in somatic and germinal tissues may be related to tissue-specific regulation of cellular metabolism.     

The Effect of 17 β-Estradiol and Estrone on Intermediary Metabolism of the Pyloric Caeca of the Asteroid Luidia clathrata (Say) Maintained under Different Nutritional Regimes

Luidia clathrata were fed either a high or low level of food and injected with estradiol, estrone, or seawater for 16 days. In individuals maintained at a low level of food, the weight of the pyloric caeca decreased in all individuals, regardless of hormone injections. Activities of all enzymes in these individuals were lower than in individuals maintained at a high level of food. In the individuals maintained at a low level of food, the activity of GPDH was higher in the pyloric caeca of individuals injected with estradiol and estrone than in those receiving seawater injections; the activity of 6-PGDH decreased in those injected with estrone. In individuals maintained at a high level of food, the activities of glucose-6-phosphate dehydrogenase (G–6–PDH) and 6-phosphogluconate dehydrogenase (6–PGDH) were higher in individuals injected with estradiol than in those receiving seawater injections. Activities of glycerol phosphate dehydrogenase (GPDH) and pyruvate kinase (PK) were not affected. A slight decrease (P<0.1) in the activity of 6–PGDH was seen with injection of estrone. These results suggest that steroids have a role in regulating cellular metabolism in the pyloric caeca, and that nutritional condition influences the response of the pyloric caeca to steroids. In addition, these results indicate that the nutritional condition of the individual directly influences enzyme activities.