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Herring appear quite unique among marine fish in having a wide range of discrete spawning seasons while using very specific spawning locations. These two aspects represent the major characteristics used to name or identify the various sub- populations. The depth and substratum of the spawning beds may vary to some extent but, for the most part, herring spawn coastally and on offshore banks, and deposit their eggs on gravel or rocks. In this respect, Baltic and Pacific herring are notably different, in that they spawn on seaweeds in the littoral or sublittoral zones.
Many recent advances in our knowledge result from technological improvements in observation and measurement. Initially, herring were assumed to spawn in schools, in response to seasonal environmental signals. Remote observation techniques, especially acoustics, do not only show schooling and vertical migration, but also demonstrate that the composition of spawning schools changes over time as new fish arrive at spawning grounds and spent fish leave Axelsen et al.
Individual females leave the spawning schools with one or more males to deposit their eggs on different sites within a particular spawning ground. Males follow closely and release ribbons of milt that are heavier than water and sink to fertilize newly spawned eggs Aneer, In Pacific herring, sperm release is triggered directly by female pheromones signalling that a female is in the process of releasing eggs Carolsfeld et al.
Egg—sperm interactions at fertilization are complex. The sperm of Pacific herring is activated only in the presence of eggs Pillai et al.
In Atlantic herring, sperm is activated on contact with seawater, although not all spermatozoa activate simultaneously: there are waves of activity that can persist over several hours Evans and Geffen, ; Geffen, Sperm motility and fertilization success vary between pairs of fish, and individual males may achieve quite different fertilization rates for eggs from different females, and vice versa.
For the most part, variations attributable to paternal contributions are not included in fishery-dynamics models. Nash et al. However, similar analyses have not yet been applied to herring. Herring serve as a good model species for studies of parental, or epigenetic, effects, because individual eggs can easily be followed throughout development and sampled just before hatching Geffen, Characteristics of the eggs and newly hatched larvae have revealed both maternal and paternal effects.
Paternal effects are evident in larva size at hatching Panagiotaki and Geffen, ; Evans and Geffen, , as well as in characteristics related to growth and metabolic potential such as protein Bang et al. Paternal effects have not yet been traced to particular characteristics of individual males, although Evans and Geffen suggested that they may relate to differences in fish size or age constituting consecutive spawning cohorts. Maternal factors are clearly demonstrated in egg size, which translates into differences in larval size at hatching Blaxter and Hempel, ; Morley, ; Morley et al. This means that the eggs and larvae are adapted to the varied seasonal conditions of temperature, light, hydrographic conditions, predator fields, and food availability.
Fish eggs and larvae are normally considered to be closely adapted to meet specific seasonal conditions match—mismatch; Cushing, The challenge has been to explain the recruitment processes in a way that matches the patterns observed in the different herring populations Cushing, ; Sinclair and Tremblay, How can herring eggs and larvae survive in all seasons? Egg sizes differ between spawning populations see below as well as between individual females Almatar and Bailey, These differences lead directly to differences in larva size at hatching Morley et al.
However, embryonic metabolism appears less subject to interactive effects between egg size and temperature in herring than in other fish species Pepin et al. Despite the population-specific differences in egg size, which may or may not be related to spawning season e. For example, herring eggs utilize the same amount of oxygen throughout development, irrespective of temperature Overnell, ; Overnell and Batty, Genetic adaptation to local conditions may have been replaced or superseded by the plasticity of eggs and yolk-sac larvae because metabolic fitness is constant over a range of temperature.
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Blaxter and Holliday described laboratory investigations from the s that examined embryonic development Brook, , ; Kupffer, and the effects of temperature on development Meyer, Early feeding studies on larvae from artificially fertilized eggs have also been reported using natural zooplankton as food Ford, ; Schach, Important advances in laboratory investigations have been made possible through Blaxter's , , , development of protocols for reliable fertilization, incubation, and rearing, as well as for keeping juvenile herring in captivity Blaxter and Holliday, Since then, studies of physiology, growth, feeding, behaviour, and responses to environmental conditions have been conducted in laboratory tanks and large mesocosms.
The results underscore how adaptable herring can be in its growth, feeding, and behaviour Munk and Kiorboe, ; Utne-Palm, At the same time, they show considerable plasticity in both physiology and form, which helps to explain the observed physical differences between populations, such as in vertebral counts and body shape Blaxter, ; Johnston et al.
Growth of herring larvae is sensitive to both temperature and feeding conditions. Unlike many other marine species, herring larvae can survive long periods of little or no growth. This ability may be part of the explanation for their survival when spawned in seasons of low productivity Johannessen et al. However, size, growth rate, and condition affect both swimming behaviour and vulnerability to predation in laboratory experiments Williams et al.
As with other species, older, well-fed herring larvae are more responsive to predator attacks and have faster escape speeds. The results from laboratory experiments have supported extensive modelling exercises testing the responses to a wide variety of environmental conditions Gallego et al. Field investigations indicate that herring larvae indeed demonstrate a wide repertoire of swimming and feeding behaviour, with a marked vertical migration component Fortier and Leggett, ; Lazzari et al. The distributions of herring larvae are influenced by water-mass features Heath et al.
The larvae of late autumn- and early winter- October—February spawning herring forage in a prey field that differs both in concentration and in composition from those of the spring and summer spawners appearing in more productive seasons. It appears an obvious question to ask what the winter-spawned larvae feed upon, but studies of their diets are few compared with those of spring- and summer-spawned larvae.
Diet preferences change in relation to larval size and ontogeny, and comparisons between seasons indicated similar preferences in the laboratory Checkley, , but significant differences in large outdoor mesocosms Gamble et al. In general, autumn-spawned larvae are smaller and consume smaller copepods such as cyclopoids Gamble et al.get link
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In the late s, otolith increment analysis was developed to estimate the age of fish larvae, and was applied with mixed success to herring studies. Townsend and Graham and Lough et al. However, Geffen found that increment counts did not correspond well with age, based on known-age larvae in tanks and in outdoor mesocosms. Otolith growth slows under poor conditions Folkvord et al.