The spatial patchiness of Pacific herring larvae

Synopsis The spatial patchiness at age was measured for Pacific herring,Clupea harengus pallasi, larvae of a small coastal inlet. Lloyd's index of patchiness decreased from 3.8 at hatching to a minimum of 1.5 at 25–30 d and then increased to 2.5 at 50 d. This two-cycle pattern resembles that measured by others for the larvae of other species of schooling pelagic fishes. It differs by being several times lower in magnitude. This is attributed to two factors: (1) a rapid reduction in patchiness from a relatively high level in the demersal egg interval to a lower level in the planktonic larval interval as larvae are released into the water column over a period of 2–4 days, and (2) foraging by larval herring for prey that are larger, more dilute, and less patchy than the prey of other, smaller pelagic fish larvae.     

Effects of larvae ontogeny, turbidity, and turbulence on prey attack rate and swimming activity of Atlantic herring larvae

The effect of turbulence, light level, and ontogeny on herring larva's attack rate and swimming activity was tested in a previous study. However, during larval seasons (spring and autumn), water clarity is frequently impaired by alga blooms, which also most probably will affect larva feeding rate. Therefore, this study was to investigate the effects of turbidity, turbulence, and ontogeny on the attack rate and swimming activity of herring larvae. By adding diatomaceous earth (DE) to the water, three turbidity levels were established: 0, 35, and 80 Jackson Turbidity Unit [JTU; which coincide with a beam attenuation (c) of 0.1, 2.5, and 4.8 m⁻¹, respectively]. An unfavourable (8×10⁻⁶ W/kg) and a favourable turbulence level (1×10⁻⁶ W/kg) were chosen based on results from the earlier study. The results show that intermediate turbidity (35 JTU) has a positive effect on the attack rate of smaller larvae (20 mm), while high turbidity (80 JTU) has a negative effect on attack rate of all tested larvae size groups. In general, attack rate was lower at the highest turbulence compared to the low level, independent of turbidity level. However, there was one exception, when turbidity was at the highest, the largest larvae (29 mm) seemed to gain from feeding in the highest turbulence level. The overall activity level was higher in the presented study than in the earlier study without turbidity. The favourable turbidity level (35 JTU) coincides with turbidity levels normally found at the equivalent depth during spring and autumn blooms in the area of where the experimental larvae originate. In addition, turbidity's effect on light absorbtion and how it influences the maximum feeding depth of the larva are discussed.     

Early life history of herring larvae in contrasting feeding environments determined by otolith microstructure analysis

Newly hatched autumn-spawned herring larvae Clupea harengus were released in two 2500-m³ outdoor mesocosms and reared over a 2-month period. Hydrographic conditions were similar in the two mesocosms, but the average plankton density was initially more than 10 times higher in mesocosm B compared to mesocosm A (>11⁻¹ v. <0.11⁻¹). Half-way through the experiment the feeding conditions reversed with three times higher average densities in mesocosm A than in mesocosm B (>31⁻¹ v. ∼11⁻¹). Herring larvae were sampled with a 0.3-m² two-chambered net twice weekly, and survivors were harvested by draining the mesocosms at the end of the experiment. Otolith growth trajectories of individual larvae were determined by relating radial otolith size with number of increments from the outer edge of the otolith (days before capture). The increment widths during the first 3 weeks after hatching, including the first-check size, were generally wider among larvae from mesocosm B (relatively good initial feeding conditions) than among those from mesocosm A (poor initial feeding conditions). The otolith growth pattern also confirmed that the surviving herring in mesocosm A belonged to the upper size range of larvae in the mesocosm after only 2–3 weeks from hatching; no such trend was found in mesocosm B. In both mesocosms the otolith size-at-age indicated that with the present sampling gear, herring larvae larger than 20–25 mm were underrepresented in the net samples. The information obtained from otolith-size-at-age is compared with other morphometric and biochemical measures of size and condition of larvae obtained throughout the experiment.     

Effects of small-scale turbulence on microalgae

Turbulence flows are characterized by their viscous dissipation rates ? and the kinematic viscosity of the fluid ν, but the effects of turbulence on organisms such as microalgae smaller than the Kolmogorov inertial-viscous length scale L_K ≡ (ν³/ε)^/14 depend on the stress τ ≡ µγ, where µ = ?ν is the dynamic viscosity, ρ is the density, and the rate-of-strain γ ≡ (ε/ν)^/12. While various workers have shown qualitatively that turbulence affects several microalgal physiological processes, these effects have not been quantified in terms of ε, τ or γ. Various microalgal groups seem to have different sensitivities to inhibition by turbulence. The relative sensitivities aregreen algae < blue-green algae < diatoms < dinoflagellates with dinoflagellates being most sensitive. We have quantified the growth sensitivity to turbulence for a red tide dinoflagellate,Gonyaulax polyedra Stein, by imposing constant γ values on cultures placed within a gap between rotating outer and fixed inner concentric cylinders. Threshold turbulence values for growth inhibition are consistent with turbulence parameters near the sea surface with light winds, suggesting turbulence may be the reason that high winds inhibit red tides. For ? > 0.18 cm²s^?3, τ > 0.04 dynes cm^?2 (0.002 N M^?2 or Pa), γ > 4.4 rad s^?1, cell numbers and chlorophyll fluorescence declined, and cells lost their longitudinal flagella and the ability to swim forward. At lower ε, τ and γ values growth rates and cell morphology were the same as in unsheared control cultures. High turbulence may affect other algae, such asSpirulina, which are commonly mass cultured.     

Effect of larval ontogeny, turbulence and light on prey attack rate and swimming activity in herring larvae

The effects of ontogeny (larval size), light and turbulence on the attack rate and swimming activity (proportion of time swimming and duration of swimming bout) of herring larvae (15-28 mm TL) have been investigated. Emphasis was put on the experimental design in order to create a set-up where the turbulence intensity distribution could be accurately measured as well as controlled in the entire experimental tank.Both larval size (ontogeny) and light had a significant positive effect on prey attack rate. Likewise, an intermediate increase in turbulence had a positive effect on prey attack rate, but this effect was dependent of light intensity and larval size.At low light (1.5 μE m² s⁻¹) intermediate turbulence increased the prey attack rate significantly for larger larvae (26 and 28 mm), while at high light (18 μE m² s⁻¹) intermediate turbulence had only a significant positive effect on the attack rate of smaller larvae 20 and 23 mm.In general, our data show a dome-shaped response of turbulence on attack rate and a U-shaped response of turbulence on swimming activity.For herring larvae >20 mm, the maximum (attack rate) and minimum (swimming activity) response of turbulence were found at intermediate turbulence intensities (energy dissipation rates between 7∗10⁻⁸ and 1∗10⁻⁶ W/kg). The highest turbulence level tested (8∗10⁻⁶ W/kg) showed only negative effects, as attack rates where at the lowest and swimming activity at the highest.Swimming activity increased with larval size or light, and decreased at intermediate turbulence. Compared to turbulent intensities under natural conditions this implies that larger herring larvae at 10 m depth have to be exposed to wind speeds of more than 17 m/s before negative effects on attack rate and swimming activity occurs.     

Effects of prey size structure and turbulence on feeding and growth of anchovy larvae

Foraging processes in plankton and planktivorous fish are constrained by relative prey and predator size and therefore, these are important variables to include in a foraging model. The distribution of prey biomass across different size classes can be characterized by a size spectrum slope. We present a foraging model for anchovy larvae including the most relevant processes such as prey encounter, capture- and pursuit success, all influenced by light, turbulence and prey characteristics. We modelled ingestion rates and specific growth rate by coupling the foraging model with an existing bioenergetic model, and performed a sensitivity analysis of prey ingestion in turbulent environments assuming either hemispherical or conical perceptive volume. Our results suggest that turbulence has no positive effect because of the low capture ability, small prey size and small visual volume for anchovy larvae. The predicted ingestion is too low to sustain the growth potential of larvae when assuming conical perceptive volume even under prey densities substantially higher than normally found in the field. Ingestion rate is sensitive to the total biomass and the slope of the prey size spectra, specifically because it determines the abundance of prey around the optimal size for the larvae. The model also suggests that small larvae benefit from a prey size structure with steep prey size-spectra slope while a large larva benefit from less steep slopes. The model can act as a link between size-spectra measurements from the field and the foraging conditions of larval anchovies.     

Metabolic recovery in herring larvae following strenuous activity

Larvae of spring spawning Clyde herring Clupea harengus L. were reared at 5 and 12° C. Metabolism following burst swimming was studied in 7-day-old larvae at their respective rearing temperatures. Escape responses were repeatedly elicited using tactile stimulation for a period of 3 min. Larval herring were hard to fatigue and still responded to tactile stimuli after 3 min. Whole larvae were freeze-quenched in liquid nitrogen, either immediately after exercise, or after periods of recovery of up to 24 h. Samples were freeze-dried and analysed for whole body creatine (Cr), phosphocreatine (PCr), ATP, ADP, AMP, lactate, glucose, and glycogen using high performance liquid chromatography and enzymatic methods. The exercise regime resulted in a marked decrease in PCr, ATP and glycogen concentrations and an increase in creatine, glucose and lactate concentrations whereas there was no significant change in either AMP or ADP concentrations. The extent of phosphagen hydrolysis (approx. 110 to 15μmol PCr g ⁻¹ dry body mass) and lactate accumulation (approx. 7 to 40 μmol lactate g⁻¹ dry body mass) over the exercise period was similar at the two temperatures, consistent with a relatively constant degree of effort. The rates of recovery of PCr and ATP were essentially the same at 5 and 12° C; returning to resting levels after approximately 30 min. Lactate and glycogen concentrations were restored 60 min after exercise at both temperatures. Maximum lactate clearance rates (1.2 μmol min ⁻¹ g ⁻¹ wet muscle mass) were an order of magnitude faster than reported for adult fish in the literature.     

On condition factor measurements in Pacific herring larvae

Condition factors (CF = mg · mm^–3 · 100) of Pacific herring larvae from Departure Bay, Canada, were determined from two larval year classes and compared for yolk sac and post-yolk sac feeding larvae. Decline in condition factors for yolk sac larvae was similar in both years. Subsequent increase of condition factors after passing the transitional phase between loss of yolk and first feeding was identical for both years. The occurrence of emaciated larvae in 1976 was attributed to poor food supply during the critical period (the time of first food uptake).     

Quantified small-scale turbulence inhibits the growth of a green alga

1. Laboratory experiments were conducted to elucidate the effect of small-scale turbulent fluid motion on the growth of laboratory cultures of the freshwater algae Scenedesmus quadricauda . Turbulent flow was generated using an oscillating-grid apparatus. The experiments were performed under the range of fluid flow conditions similar to those occurring in nature. The only growth limiting factor was the effect of small-scale fluid motion; all other environmental factors, such as light, temperature and nutrients, were kept constant.
2. Growth of Scenedesmus quadricauda , measured in terms of chlorophyll a concentration, was inhibited when the level of turbulence in the water column was increased. Algal growth was maximum in a quiescent fluid. The inhibitory effect of fluid motion was observed independently of flow regime (laminar, transitional, turbulent) in the water column.
3. Cell destruction and aggregation of dead and living cells of algae were observed in a turbulent flow. High shear rates, estimated from the dissipation of turbulent kinetic energy, caused the cell destruction, algal collision and agglomeration of algae. Data on Scenedesmus responses to small-scale fluid motion will enhance and broaden our ability to develop predictive multispecies models for freshwater phytoplankton communities.     

Effects of small-scale turbulence on feeding rate and gross-growth efficiency of three Acartia species (Copepoda: Calanoida)

Experiments designed to test the effects of small-scale turbulenceon the feeding rates and gross-growth efficiency of calanoidcopepods have been performed within a wide range of controlledfood concentrations. Turbulence significantly enhanced feedingrates only at food concentrations lower than the ingestion saturatinglevel. Gross-growth efficiency (the quotient carbon-egg production/carbon-foodingested) of the different Acartia species studied showed differentpatterns of response to turbulence, in agreement with the hydrodynamiccharacteristics of their habitat. Furthermore, experiments conductedon Acartia clausi at two different intensities of turbulenceindicate a shift in the response, with enhancement of feedingat low intensities of turbulence and negative interference athigher intensities.     

Extreme helmet formation in Daphnia cucullata induced by small-scale turbulence

Although Daphnia cucullata is used as a textbook example forcyclomorphosis, distinct helmet development, as shown in thefield, has not been demonstrated in the laboratory until now.We show for the first time that small-scale turbulence is ableto induce the maximum response of morphological plasticity inD. cucullata. Helmet elongation reached magnitudes as extremeas observed in the field. Different modes of generating small-scaleturbulence caused different levels of helmet formation. Small-scaleturbulence alone may not be responsible for cyclomorphosis innature because field data show that two nearby lakes with similarmorphometry differ in cyclomorphosis patterns, while laboratoryexperiments show that there is no difference in the abilityto form helmets in the clones of each lake. Although helmetformation in D. cucullata is inducible with predator kairomones,helmet elongation is not as strong as that induced by turbulence.We discuss the possible role of helmets under turbulent conditions.     

The importance of managing within-species diversity in cod and herring fisheries of the north-western Atlantic

The spatial structure of several exploited species within and among known populations of the Northwest Atlantic is reviewed using past and current knowledge. For each species, the coherence of existing management units with the spatial scale of population dynamics (populations, sub-populations, and spawning components) is summarized. The implications of this match or mismatch of spatial scales in the maintenance of biodiversity within and among populations are also discussed.     

The role of escape reactions in determining the size distribution of prey captured by herring larvae

Synopsis Fish larvae are selective planktivores, and size is an important factor in prey selection. However, for herring larvae, a selection model based solely on the principle of optimising calorific gain per unit energy expenditure consistently overestimates the mean size of ingested prey. Most such models ignore the escape capabilities and shape of the prey, and the potential role of escape on capture success. In this paper, a static foraging model incorporating both selection and escape is described and tested against published data on the composition of herring larvae stomach contents. The results indicate that prey escape is a major factor structuring the diet composition of the larvae.     

Susceptibility of Atlantic herring and plaice larvae to predation by juvenile cod and herring at two constant temperatures

Predation encounters were staged in the laboratory to compare prey responsiveness, predator error rate, and predator capture success for juvenile cod Gadus morhua (a suction feeder) and herring Clupea harengus (a biting predator) preying on herring and plaice Pleuronectes platessa larvae. Trials were conducted at near natural temperature extremes for the larvae (8 and 13°C) to assess the importance of water temperature to the interaction. Herring larvae were significantly more responsive to attacks by herring than were plaice larvae (5·7 vs 0'0%). The two prey species were equally responsive to attacks by cod (2–6 vs 10%). Cod caught 91% of herring larvae attacked and juvenile herring caught 87%. Cod were successful in 96% of attacks on plaice, but juvenile herring caught significantly fewer (83%) plaice larvae. For each predator species, capture success did not vary significantly with prey species. Overall capture success for herring was significantly lower than that for cod. Responsiveness of herring larvae to attacks by juvenile herring increased with temperature, but predator error rate and capture success were not altered by water temperature.     

The larvae of some blowflies of medical and veterinary importance

Diagnostic features are described as a series of couplets that enable separation of the third instar larvae of the following pairs of closely related forms of blowflies of medical and veterinary importance: Chrysomya chloropyga (Wiedemann) and Ch.putoria (Wiedemann), Chrysomya albiceps (Wiedemann) and Ch.rufifacies (Macquart), Cochliomyia hominivorax (Coquerel) and Co.macellaria (Fabricius), Lucilia sericata (Mergen) and L. cuprina (Wiedemann), Calliphora augur (Fabricius) and C. stygia (Fabricius).     

Influence of temperature on evasive responses of Atlantic herring larvae attacked by yearling herring, Clupea harengus L.

Predation encounters were staged in the laboratory between yearling herring, Clupea harengus L., 66 to 104 mm t.l. , and herring larvae, 8 to 30 mm t.l. ., at 8,11 and 14 o C. Video records were used to quantify prey behaviour. Prey responsiveness, reactive distance, response latency, and apparent looming threshold were not affected by temperature. Response speeds increased with temperature. Predator error rate and capture success showed no consistent thermal effects. Although the experiments could not fully evaluate the influence of temperature on the predators, results suggest that the predator's performance largely governs the outcome of an attack on a larva and that higher temperatures favour the predator by increasing the frequency of its encounter with prey.     

Diel differences in avoidance of three vertical profile sampling gears by herring larvae

Three plankton sampling systems [pump, Longhurst-Hardy PlanktonRecorder (LHPR) and MOCNESS] were used to obtain vertical profilesof herring larvae off the north-east coast of England in October1986 during the International Herring Larval Survey (IHLS).A factorial sampling design allowed a comparison of the numberof herring larvae caught by depth (four levels), time of day(three levels) and gear (three levels). The size compositionof herring larvae taken in the different gears has also beenexamined. Catches taken at night were greater than those takenduring the day throughout the water column, particularly withlarger larvae. This indicates that visual avoidance of all gearsis an important factor, which increases as the larvae grow bigger.LHPR night catches were greater than in the MOCNESS and fargreater than in the pump, which indicates that avoidance isalso due to senses other than vision. Examination of the 1981–1986International Herring Larvae Survey catch data showed that night-timecatches were similar to daytime catches for herring larvae <10mm long, but night time catches caught progressively more asthe length of the larvae increased to 24 mm.     

The importance of the behaviour of young larvae in sorghum resistance to Chilo partellus

Management of spotted stem borer, Chilo partellus (Swinhoe) by the use of resistant cultivars is being developed at ICRISAT. Different cultivars show resistance based on several mechanisms, one of which affects the establishment of first instar larvae in the plant whorl. However, it has been found that low establishment is a factor associated with resistance only in some cultivars and not in others. A number of physical and chemical plant factors are associated with preferential establishment of the young larvae in the leaf whorl. Among the physical characteristics, erect leaves and curled leaf bases are involved in larval establishment. Detailed chemical analysis of surface extracts showed that the concentration of one compound in the wax is related to resistance at this stage and assessment of physical and chemical plant characters has been used to predict resistance.

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Résumé L'Institut International de Recherches sur les Cultures en Régions Semi-Arides (ICRISAT) étudie la protection contre C. partellus Swinhoe par l'utilisation de cultivars résistants. Différents cultivars présentent une résistance à différents stades de croissance de la plante, et à différents stades de l'insecte. La résistance du sorgho à C. partellus dépend de plusieurs mécanismes, l'un d'eux conditionnant l'installation des chenilles du premier stade dans le verticille de la plante. Cependant, la lenteur de l'installation d'est un facteur lié à la résistance que chez quelques cultivars. Plusiers facteurs physiques et chimiques sont associés à l'installation préférentielle des jeunes chenilles dans le verticille foliare. Parmi les caractères physiques, le port érigé des feuilles, la présence de poches à leur base et les cires superficielles sont impliqués dans l'installation des chenilles. Une analyse chimique détaillée d'extraits des structures superficielles a montré que la concentration d'une substance, le C32 marqueur de la lignée résistante IS 2205, était 2 fois plus faible que dans les lignées sensibles IS 1151 et CSH 1.

     

The importance of 'memory' in statistical models for animal feeding behaviour

We investigate models for animal feeding behaviour, with the aim of improving understanding of how animals organise their behaviour in the short term. We consider three classes of model: hidden Markov, latent Gaussian and semi-Markov. Each can predict the typical 'clustered' feeding behaviour that is generally observed, however they differ in the extent to which 'memory' of previous behaviour is allowed to affect future behaviour. The hidden Markov model has 'lack of memory', the current behavioural state being dependent on the previous state only. The latent Gaussian model assumes feeding/non-feeding periods to occur by the thresholding of an underlying continuous variable, thereby incorporating some 'short-term memory'. The semi-Markov model, by taking into account the duration of time spent in the previous state, can be said to incorporate 'longer-term memory'. We fit each of these models to a dataset of cow feeding behaviour. We find the semi-Markov model (longer-term memory) to have the best fit to the data and the hidden Markov model (lack of memory) the worst. We argue that in view of effects of satiety on short-term feeding behaviour of animal species in general, biologically suitable models should allow 'memory' to play a role. We conclude that our findings are equally relevant for the analysis of other types of short-term behaviour that are governed by satiety-like principles.