Effect of temperature on the escape responses of larval herring,Clupea harengus

Herring (Clupea harengus L.) larvae from spring and autumn spawning stocks were reared at different constant temperatures from 5° to 17 °C. At equivalent developmental stages, the spring larvae were longer than the autumn larvae and the larvae reared at low temperatures were longer than those reared at high temperatures. At hatching and at the end of the yolk-sac stage, the larvae were induced, by a probe, to make C-start escape responses, which were recorded and analysed using a high-speed video recording at 400 frames s^-1. The response was rapid and of short duration. The tailbeat frequency and swimming speed were measured during the burst of swimming following the C-start at different test temperatures and in larvae with different temperature histories. The tail-beat frequency was strongly temperature-dependent, rising from 19 Hz at 5 °C to 37 Hz at 17 °C with no effect of temperature history, season or developmental stage. The burst-swimming speed ranged at hatching from 75 to 90 mm s^-1 at 5 °C to 110 to 160 mm s^-1 at 17 °C and at yolk resorption from 90–115 mm s^-1 at 5 °C to 175–190 mm s^-1 at 17 °C. The longer, spring-spawned larvae swam faster than the shorter autumn-spawned larvae. When the swimming speeds were expressed as body lengths (L) s^-1, these differences disappeared. Larvae swam from 7–9 L s^-1 at 5 °C to 15–20 L s^-1 at 17 °C at hatching, and from 8–9 L s^-1 at 5 °C to 15–17 L s^-1 at 17 °C at yolk resorption. There was, however, a significantly faster specific swimming speed by the larvae reared at 12 °C in spring 1991.Honorary Research Fellow of the Scottish Association for Marine ScienceUnfortunately, Karen Fretwell was drowned in an accident on 9 January 1993     

Light intensity and the feeding behaviour of herring,Clupea harengus

An infra-red sensitive video-recording technique was used to study the effects of darkness and light intensities from 0.0001 to 270 photopic lx on the feeding behaviour of herring (Clupea harengus L.). When offered natural zooplankton, consisting of a mixture ofCalanus finmarchicus, Euchaeta norvegica, Oithona similis, Balanus sp. nauplii, and crustacean nauplii as prey, the fish fed by biting (snapping) at light intensities above a threshold of 0.001 lx and were size-selective, taking the larger organisms first. When fed on pure cultures of CaliforniaArtemia sp. nauplii (San Francisco Bay brand), the threshold light intensity was 0.01 lx. Swimming speed increased with increasing light intensity when the fish were actively feeding by biting. When the fish were filter-feeding on high densities ofArtemia sp. nauplii in the light, they continued to school and swimming speed was not related to light intensity.     

A laboratory study of predation by Aurelia aurita on larval herring (Clupea harengus): Experimental observations compared with model predictions

Predation by the medusa Aurelia aurita L. on early first-feeding stage larvae of the herring clupea harengus L. was studied in the laboratory. The medusae were captured in Loch Etive, Scotland. Herring larvae were reared from the extificially fertilized eggs of spawning Clyde herring caught in March, 1982. Swimming speeds, volume searched”, capture efficiency and predation rates increased as medusa size increased. Predation rates on fish larvae increased with prey density, but appeared to approach a maximum at high prey densities; in 1 h experiments, a maximum rate of predation of 6.64 larvae h^-1 was estimated by fitting an Ivlev function. A model to predict predation rates was constructed from swimming speeds, sizes and densities of medusae and larvae, and capture efficiency. The rates of predation predicted from the model fell within the range of experimental data, but tended to underestimate rates and did not account for saturation of medusae. Swimming patterns of medusae changed after prey capture: (a) before capture, encounter rates were low and medusae were relatively less active; (b) after capture of 1 larva, encounter rates doubled, with the stimulated medusae exhibiting increased activity and an aftered “searching” path; and (c) after capture of many larvae, swimming speeds and encounter rates of medusae decreased.     

Osmotic properties of eggs of the herring Clupea harengus

Yolk osmolality of developing eggs of the herring Clupea harengus L. is strongly hypoosmotic to seawater: about 440 mOsm during the first week of development decreasing to 360 mOsm before hatching. The perivitelline fluid (PVF) of the eggs is isoosmotic to the ambient medium. The PVF equilibrates within 10 min to changes in the ambient seawater. The content of Na⁺, K⁺, Cl^- and free amino acids amount to 26, 52, 48, and 54 n mol egg^-1, respectively, on Days 1 to 3 after fertilization, increasing to 63, 69, 80, and 79 n mol egg^-1, respectively, prior to hatching (Days 18–20). The apparently conflicting findings of a decreasing yolk osmolality and a simultaneous increase in the amount of egg solutes are at present unexplained.     

Variations in the content of trypsin and trysinogen in larval herring (Clupea harengus) digesting copepod nauplii

Trypsin and its proform trypsinogen were quantified by radioimmunoassay in herring (Clupea harengus L.) larvae subjected to different prey densities. During the first weeks of larval life, the enzyme content fluctuated in a threephased pattern. Yolk resorption (Phase 1) was characterized by an increase in enzyme. During the first few days after yolk resorption (Phase 2), there was a sharp decline in enzyme. Older larvae (Phase 3) exhibited a second period of intensive enzyme synthesis. Amounts of trypsin in intestines of feeding larvae were analysed. At first feeding, a basal level of gut enzyme of approximately 30ng was recorded, and the amount of additional enzyme secreted from the pancreatic tissue into the intestine appeared to be dependent upon the numbers of prey items ingested. The enzyme-substrate ratio in the intestine was approximately 1 to 4. Prey availability affected amount of trypsinogen. Larvae experiencing a high prey density had an approximately two-fold higher specific enzyme content in Phase 2 compared to larvae exposed to a low prey density. A proposed nutritional strategy for first feeding herring larvae is discussed.     

Growth and mortality in young larval herring (Clupea harengus); effects of repetitive changes in food availability

Following yolk resorption, laboratory-reared larval Baltic herring (Clupea harengus L.) were exposed to two sequences of food restriction for 5 d and re-alimentation for 10 d. Comparisons regarding larval growth (standard length and content of water-soluble protein), mortality and content of the sum of trypsin and trypsinogen were made with larvae at a continuous high ration. Larvae exposed to varying prey abundance grew less in length than the control, and during the second high-ration period (Day 22 to 32) growth in length ceased. From the first low-ration period onwards, the content of water-soluble protein in these larvae was lower than that of the control larvae, and the survival rate of the low-high ration group was 59% compared to 77% in the larvae at a continuous high ration. In contrast, the effects of varying food availability were minor on larval content of trypsin and trypsinogen. Results are compared with previous findings in larval Clyde herring, and the effects of larval stock and timing and duration of food restriction on larval growth performance are discussed.     

Effects of a transitory,low food supply in the early life of larval herring (Clupea harengus) on mortality,growth and digestive capacity

In order to assess possible effects of a transitory, low food supply on later development, three groups of Clyde herring larvae (Clupea harengus L.) were exposed in 1989 to different feeding regimes immediately after yolk resorption. Group 1 received a high daily ration of 80 copepods larvae^–1 for 31 d, Group 2 a low daily ration of 15 copepods larva^–1 for 10 d followed by a high ration (80 copepods larva^–1) for 21 d and Group 3 a low ration of 15 to 20 copepods larva^–1 for 31 d. After 31 d of feeding, digestive capacity, expressed as the sum of trypsin and trypsinogen, was markedly reduced in Group 2 compared to Group 1, while Group 3 had an even lower digestive capacity. After the switch from low to high ration Group 2 exhibited compensatory growth and caught up with Group 1 both in standard length and content of soluble protein. Group 3 had the lowest growth rates. Mortality was equal in Groups 1 and 2, while Group 3 showed an excess mortality of 40% of the start population. Although Group 2 larvae had caught up with Group 1 in growth at the end of the study, content of trypsin and trypsinogen in Group 2 was only half of that found in Group 1. Thus, comparing effects of a short period of food limitation on future growth, mortality and content of digestive enzymes, the study indicates content of trypsin and trypsinogen to be the most sensitive variable for detection of food limitation in the early stages of exogenous feeding.     

Effects of sulfuric water pollutants on fertilization,embryonic development and larvae of the herring,Clupea harengus

Using herring (Clupea harengus) eggs and larvae from Baltic spring spawners, the biological effects of sulfuric pollutants (largely FeSO₄ and H₂SO₄) which are scheduled to be released daily into the North Sea in large quantities, have been tested. Dilutions (1:8000, 1:16000, 1:24000 and 1:32000) of the sulfuric pollutants were used as test media. Throughout all experiments water temperature was maintained at 8.0°±0.2°C, salinity near 16.5; eggs were artificially fertilized 1 or 6 to 7 h after catching the parent individuals and, attached to glass plates, incubated in 1 l aerated containers. In all test media brownish precipitates, which resulted from diluting the sulfuric pollutants with 50% sea water, tend to adhere to the egg surfaces, thereby possibly interfering with gaseous and other exchanges between egg and surrounding medium. Under conditions of maximum test medium effectiveness, percentages of successful fertilization and of egg survival are considerably reduced; diameter of fertilized eggs remains smaller; embryonic growth rate is retarded while the heart frequency tends to increase (indicating physiological stress); duration of incubation is shortened: percentage of successful hatching decreases and structural abnormalities of freshly hatched larvae increases. Dilutions down to 1:24000 and 1:32000 have rather limited effects. Exposure of 1 to 3 day old, healthy herring larvae to the 4 test media leads to failure of performing prey catching manoeuvres in 1:32000 and 1:24000, to impaired locomotory performances in 1:16000, and practically to paralysis, shrinkage, permanently bent bodies and death within a few days in 1:8000. Although the present study needs deepening through further and more detailed experiments, it can be said that the pollutants under consideration represent a danger to herring eggs and larvae at least up to a dilution of 1:32000.     

Biological effects of oil on early development of the Baltic herring Clupea harengus membras

The effects of petroleum hydrocarbons from two crude oils and one fuel oil (No. 1) were studied on the ontogenic development of the Baltic herring Clupea harengus membras L. Herring eggs exposed to water-soluble fractions of the oils at the time of fertilization showed no decrease in numbers of fertilized eggs compared to eggs exposed 6 or 72 h after fertilization. During embryongenesis, treatment with 3.1 to 8.9 ppm or 3.3 to 11.9 ppm total oil hydrocarbons from light fuel oil and the two crude oils respectively, gave rise to alterations in embryonic activity, decreased heart rate, and premature or delayed hatching. Although many larvae hatched from eggs exposed to contaminated water (3.1 to 11.9 ppm total oil hydrocarbons), the majority of the (70 to 100%) were malformed or dead 1 day after hatching. Exposure of eggs to 5.4–5.8 ppm total oil hydrocarbons resulted in significantly (P<0.001) decreased lengths of the larvae. Increased temperature (from 9° to 14°C) aggravated the effects of the oils. The results are discussed in relation to the potential effects of oil spills and chronic oil pollution on fish eggs and larvae in the Baltic Sea.     

Male characteristics, sperm traits, and reproductive success in winter-spawning Celtic Sea Atlantic herring, Clupea harengus

The relationship between sperm characteristics and reproductive success was examined in male herring, Clupea harengus L. Males were categorised as being first-time or repeat spawners on the basis of their age; they were also grouped according to whether their sperm were immediately active and exhibited forward motion on contact with seawater (FM) or had little or only vibratory motion (VM). Unlike the Pacific herring C. pallasii Valencienes, Atlantic herring sperm is usually motile on contact with seawater. The age, weight and gonadosomatic index (testes mass as a percentage of somatic mass = GSI) were measured and used as characteristics for individual fish. Sperm traits measured were (1) adenosine triphosphate (ATP) concentration, (2) sperm count, (3) duration of sperm motility. Reproductive success for each male was estimated from the fertilisation rate and from the length of larvae at hatching. Fertilisation rates for all fish were generally >80%. The ATP concentration of non-activated spermatozoa was negatively correlated with fertilisation rate. Among repeat spawners, fish with higher GSIs produced larvae that were larger at hatching. Although VM sperm fertilised eggs at rates equivalent to fertilisation by FM sperm, the larvae produced by VM sperm were significantly smaller at hatching. Larval length tended to increase in parallel with the duration of sperm motility, but the relationship was not significant in these tests. The results did not indicate any age or size pattern to spawning readiness in male herring. Sperm that are not yet ready to be shed are not fully motile on contact with seawater, but are still capable of fertilising eggs that hatch successfully. There is likely to be a progression of males which come into spawning readiness within a spawning shoal; therefore it is possible that paternal influences would result in a progressive decrease in larval size over the spawning period in winter-spawning Celtic Sea herring. Received: 22 November 1997 / Accepted: 8 June 1998     

Escape behaviour of solitary herring (Clupea harengus ) and comparisons with schooling individuals

The escape behaviour of solitary herring (Clupea harengus L.) startled by a sound stimulus was observed by means of high-speed video-filming. The results were compared with data from a previous study on the escape behaviour of schooling herring. Escape responses were divided into “away responses” and “towards responses” according to the orientation of the C-bend of the body relative to the stimulus. The proportion of away responses was smaller for solitary than for schooling herring. In solitary herring, the subsequent escape trajectories of fish making initial away responses showed a bimodal pattern of distribution, with modes at 130 and 180° from the stimulus. Trajectories following towards responses, however, were mainly within the semicircle directed at the stimulus, and their pattern of distribution differed from that of away responses. This result contrasts with observations on schooling herring, whose trajectories following both initial away and towards responses are directed away from the stimulus. In addition, we measured the response latency, defined as the interval of time between stimulus presentation and the first detectable movement of the fish. Solitary herring showed a higher proportion of short-latency responses (latency <50 ms) than schooling herring. Different behaviours appear to be exhibited by herring depending on whether they are solitary or within a school. We hypothesize that schooling may raise the threshold for initiation of fast escape responses, giving longer latencies and slower responses which are more appropriate in their directionality and reduce the possibility of collisions with neighbours. In addition, we suggest that schooling behaviour enhances the directionality and co-ordination of the escape response of the whole school, possibly increasing the probability of surviving a predator attack. Received: 2 October 1996 / Accepted: 17 October 1996     

RNA:DNA ratios and growth of herring (Clupea harengus) larvae reared in mesocosms

Autumn-spawned North Sea herring larvae (Clupea harengus L.) were released in two outdoor mesocosms of 2500 m³ (A) and 4000 m³ (B). The mesocosms were monitored for temperature, salinity, oxygen, chlorophyll a, zooplankton and herring larvae abundance. The density of suitable prey for first feeding larvae (mainly copepod nauplii) was initially low in Mesocosm A (<0.11^-1) compared to in Mesocosm B (>11^-1). Half-way through the experiment the situation was reversed, with higher densities of prey in Mesocosm A (>31^-1) as compared to Mesocosm B (～11^-1). The average temperature declined steadily in both mesocosms from 18°C at release to 11–12°C by the end of the experiment 60 d later. The RNA:DNA values of individual herring larvae were related to protein growth rates and temperature adjusted according to Buckley (1984). A corresponding DNA growth index (Gdi) was given as: Gdi=0.68 TEMP+3.05 RNA:DNA-9.92. The RNA:DNA based growth indices were significantly correlated with other somatic growth estimates. The average estimated protein growth rate in the two mesocosms followed the same temporal pattern as the somatic growth rate, but with a lag of 2 d or more. Residual analysis of the regression of ln RNA versus ln DNA also showed the same temporal pattern as the RNA:DNA ratios, but the shift in condition as estimated by this method occurred more in synchrony with the other somatic growth measures. Larvae in Mesocosm A had RNA:DNA values similar to the starvation control kept in the laboratory the first days after release, confirming that larvae in Mesocosm A initially were in poor nutritional condition. On the other hand, the majority of the herring from Mesocosm B were characterised as starving or in poor nutritional condition towards the end of the experiment. The assessment of growth and nutritional condition were in accordance with independent survival estimates which suggested that the majority of the total mortality occurred during the first 15 d in Mesocosm A and there-after in Mesocosm B.     

Developmental changes in the composition of myofibrillar proteins in the swimming muscles of Atlantic herring,Clupea harengus

Changes in myofibrillar protein composition during development have been investigated in the swimming muscles of the Atlantic herring Clupea harengus L. using a range of electrophoretic techniques. The main muscle-fibre type of larvae, and the fast- and slow-muscle fibres of adult fish were found to contain distinct isoforms of myosin heavy chain (MHC) and myosin light chain 2 (LC2). Larval LC2 was present as a minor component of adult fast-muscle myosin. In contrast, larval and adult fast-muscle myosin appeared to contain identical alkali light chains. Tropomyosin and troponin C were also identical in larval and in adult fast-muscle. All three muscle-fibre types contained unique isoforms of troponin T (TNT) and troponin I (TNI). Larval muscle had multiple isoforms of TNT, some of which may correspond to embryonic forms. It was concluded that although the main muscle-fibre type in larvae shares some myofibrillar proteins with adult fast muscle, it also contains characteristic isoforms of MHC, TNI, TNT and LC2 and therefore represents a distinct fibre type. The particular combination of myofibrillar proteins present at any developmental stage was found to be dependent on the rearing temperature. For example, a higher proportion of embryonic TNT isoforms were present at hatching in larvae reared at 5°C than at either 10 or 15°C. Over a period of 7 d, there was a gradual reduction in the number of TNT isoforms, but the pattern in 5°C larvae after 7 d still did not resemble that in 1 d-old larvae reared at 15°C.     

Factors influencing predation of Hyperoche medusarum (Hyperiida: Amphipoda) on larvae of the Pacific herring Clupea harengus pallasi

Predation of different-sized Hyperoche medusarum (Hyperiida: Amphipoda) on larvae of the Pacific herring Clupea harengus pallasi was studied in the laboratory. The attacking rate of H. medusarum was a function of herring larvae size as well as size of the predator, and varied from 0.15 to 0.95 larvae attacked h^-1 per hyperiid. In the range of 7.55 to 16.05 mm total larval length, vulnerability to predation was highest for 13.3 and 13.7 mm larvae. Large hyperiids swam faster and covered a wider area during searching and were more effective predators than small ones. Predation seemed to be influenced by light, and its intensity was dependent on the duration of previous food deprivation of the hyperiid.This study was sponsored by the International Bureau of the Gesellschaft für Kernenergiever-wertung in Schiffbau und Schiffahrt in connection with the German Canadian agreement on scientific and technical cooperation.     

Trypsin content in intestines of herring larvae,Clupea harengus,ingesting inert polystyrene spheres or live crustacea prey

In 1986, at the Danish Institute of Fisheries and Marine Research, Denmark, Clupea harengus L. larvae from three different herring stocks were offered either non-biodegradable polystyrene spheres, nauplii and copepodites of Acartia tonsa or Artemia ssp. nauplii. Ingestion of polystyrene spheres induced trypsin secretion to a higher level than in non-feeding fish. Larvae ingesting live food of the same width as the polystyrene spheres exhibited the highest trypsin content in the intestines. Mechanisms responsible for the regulation of pancreatic enzyme secretion are discussed.     

Temperature influences muscle differentiation and the relative timing of organogenesis in herring (Clupea harengus) larvae

Eggs from spring spawning stocks of herring (Clupea harengus L.) were fertilized and reared at either 5, 8 or 12°C in 1991 and 1992. The differentiation of myotomal muscle fibres was investigated in relation to the development of other organs and tissues using light and electron microscopy. The gut, notochord, eyes and haemocoel appeared at the same relative point in development between fertilization and hatching at all temperatures. In contrast, the formation of the spinal cord, pronephros, pectoral fin buds and muscle fibres was relatively retarded at 5°C compared with 8 and 12°C. Myogenesis in the presumptive inner muscle mass occurred after 12 to 16 d at 5°C, 7 to 10 d at 8°C and 3.5 to 6 d at 12°C. Myoblasts aligned in orderly rows running from myosept to myosept prior to fusion to form myotubes. Actin and myosin filaments were synthesised throughout the cytoplasm in associated with presumptive Z-lines at the periphery of myotubes and immature muscle fibres. Differentiation of the superficial and inner muscle fibres types of larvae occurred at around the same time. Following this initial period of myogenesis, the number of myotomal muscle fibres remained constant until after hatching, so that increases in muscle bulk in the late embryo were entirely due to fibre hypertrophy. At hatching, the number of superficial muscle fibres present in myotomes just posterior to the yolk-sac was significantly less at 5°C (108±12) than at either 8°C (132±10) or 12°C (140±10) (mean±SD, 12 fish/temperature). In contrast, there were around 280 inner muscle fibres/myotome, comprising 90% of the trunk cross-sectional area, at all three temperatures. Myofibrillargenesis occurred relatively slowly at low temperatures, so that the volume density of myofibrils in the inner muscle fibres of larvae at hatching was significantly less at 5°C (39.2±9.0) than at either 8°C (49.6±8.8) or 12°C (50.2±9.8) (mean ±SD, 20 fibres/temperature from total of 5 fish). Undifferentiated myoblasts remained at hatching to form a population of presumptive myosatellite cells. The number of presumptive myosatellite cells per mm² cross-sectional area of muscle fibre was more than two times higher at 8°C (1493±335) than at either 5°C (478±102) or 12°C (924±233) (mean±SD, 5 fish/temperature). The results suggest that temperature can influence the commitment of myoblasts to differentiation at a critical stage in embryogenesis, thereby providing a potential mechanism for influencing future growth characteristics. Correspondence to: I.A. Johnston at Gatty Marine Laboratory     

Effect of short-term variations in light intensity on photosynthesis of a marine phytoplankter: A laboratory simulation study

Parameters of photosynthesis and growth were measured for a marine diatom (Lauderia borealis) grown in axenic continuous culture under three different light regimes: constant, simulated diurnal variation, and fluctuating. The light fluctuations were systematic increases and decreases in light intensity superimposed on the diurnal regime. In the first two regimes, a morning maximum and an afternoon depression in photosynthesis were observed. In the fluctuating light regime, the afternoon depression was less pronounced and the morning maximum was enhanced. The results may be explained by postulating a time-dependent value for the light-saturated rate of photosynthesis. Light utilization [mmol O₂ cell^-1 (E m^-2)^-1] was the same for the diurnally varying and fluctuating regimes, despite the fact that the peak light intensity in the fluctuating regime was double that of the diurnally varying regime.     

Quantification of trypsin with a radioimmunoassay in herring larvae (Clupea harengus) compared with a highly sensitive fluorescence technique to determine tryptic enzyme activity

Enzymatic activity and quantity of the protease trypsin were measured in individual herring larvae (Clupea harengus L.). The enzymatic activity assay was done using a fluorescence technique, and a radioimmunoassay was used for quantification of trypsin. The results are compared and the differences between the techniques discussed. Both methods gave similar results, as high or low values in trypsin quantity were reflected in high or low values of tryptic activity. Quantity and activity were linearly and positively correlated, but small differences between methods were found at the lowest detection limits. Both techniques reflect high variability between individual larvae.     

Stimulatory effect of ingested protein and/or free amino acids on the secretion of the gastro-endocrine hormone cholecystokinin and on tryptic activity, in early-feeding herring larvae, Clupea harengus

In the larvae of many marine teleosts, the stomach is absent until they approach or attain metamorphosis. Consequently, the formation of chyme containing specific free amino acids from the gastric digestion of protein, which are believed to be signals initiating the release of the digestive hormone cholecystokinin (CCK), is lacking. CCK, when secreted into the blood circulation from specialized intestinal cells, stimulates gallbladder motility and is a key factor causing the release of pancreatic digestive enzymes into the gut lumen. Using first-feeding Atlantic herring larvae (Clupea harengus) as a model, the aim of the present study was to determine if a CCK response together with tryptic activity could be elicited in larvae ingesting dietary protein and/or FAA. Larvae were tube fed single lamellar liposome vesicles (SLV) containing: (1) physiological saline (PS), (2) bovine serum albumin (BSA), (3) specific free amino acids (FAA), or (4) a ratio (1:1) of BSA and FAA. The CCK and trypsin levels were then assayed (radio-immunoassay) at 0, 15, 60 and 120 min after tube feeding. A marked CCK response was elicited in all treatments compared to the PS control at 15 and 30 min and was significant (p<0.05) at 120 min after tube feeding. Larvae tube fed the FAA treatment exhibited CCK levels that increased linearly from 1.6 to 5.6 fmol mg^-1 dry weight (DW) after 2 h of digestion, although this response was below the BSA and BSA:FAA treatments. The BSA and BSA:FAA treatments, after 15 min of digestion, showed a rapid CCK increase, over the PS and the FAA liposome treatments, to 8.1 and 5.4 fmol mg^-1 DW, respectively. At the end of the assay, BSA and BSA:FAA demonstrated similar levels (10.2 and 9.2 fmol mg^-1 DW, respectively). Larvae tube fed the PS control or the FAA liposome treatment did not demonstrate any appreciable increase in tryptic activity during the 2 h digestion period (0.03-0.071 and 0.03-0.048 mU mg^-1 DW, respectively). In contrast, the BSA:FAA treatment increased from 0.03-0.148 mU mg^-1 DW 1 h after feeding, which was significantly (p<0.05) higher than the PS and FAA liposomes, and then decreased markedly (0.085 mU mg^-1 DW) after 2 h of digestion. The larvae tube fed BSA liposomes, however, demonstrated steadily increasing tryptic activity throughout the sampling period, attaining 0.255 mU mg^-1 DW after 2 h, which was significantly (p<0.05) more than all the other treatments. The results showed that ingested liposomes containing FAA or the protein BSA or a combination of these two nutrients effectively stimulated CCK production in first-feeding herring larvae. In contrast, liposomes containing only physiological saline did not elicit a CCK response. In addition, liposomes containing BSA stimulated tryptic activity in herring larvae, which was not observed in fish fed liposomes that included only FAA or PS. This suggests that a suitable protein substrate is required to regulate protein digestion.     

In-situ observations of Baltic herring (Clupea harengus membras) spawning behaviour in the Askö-Landsort area,northern Baltic proper

In a Baltic herring (Clupea harengus membras L.) spawning ground study, carried out in summer 1982 in the northern Baltic proper, spawning herring were encountered on two line transects on June 14. The spawning behaviour of minor groups as well as of a large school was studied in situ by four divers for approximately three hours. No behavioural differences or systematic coordinated behaviour between the sexes could be detected. Most of the spawning took place in a limited depth interval, 0.5 to 4 m, and Ceramium tenuicorne was the substrate most used.