Digestive system functionality is a key process linked to larval recruitment and survival. However, little is known about organ development and enzyme maturation of the digestive system of North Sea Atlantic herring (Clupea harengus). In this study, herring larvae were reared at 13 °C from hatching to 69 day post hatch, covering four developmental stages: (1) yolk sac (8-9 mm), (2) pre-flexion (9-14 mm), (3) flexion (12-18 mm) and (4) post-flexion stages (15-30 mm). Combined histological (semi-quantitative scoring) and enzyme analyses (pancreatic and intestinal) showed that developmental stages are strongly linked to physiological changes. The larvae lack a functional stomach and use the intestine as the primary site of digestion which is mainly supported by pancreatic enzyme activity. The intestine acquired adult enzymatic digestive features with a functional brush border at the end of the flexion stage and pyloric ceca started to develop during the post-flexion stage. The transition from pre-flexion to flexion stage and the end of the post-flexion stage are energetically taxing periods as indicated by a reduced number and size of liver vacuoles. Based on these findings, we consider these moments as critical periods, where herring larvae could be dramatically affected by suboptimal feeding conditions in the field. This implies that pre-flexion stage larvae with low or no liver reserves may not be able to proceed to the next developmental stage. Hence, the level of energy storage in first-feeding larvae needs to be examined for its use as a field indicator of survival and development.

Condition indices aim to evaluate the physiological status of fish larvae by estimating both the level of starvation and potential of survival. Histological indices reveal direct effects of starvation whereas biochemical indices such as lipid classes or RNA:DNA ratios are used as proxies of condition, giving information on the amount of energy reserves and growth rate, respectively. We combined these three indices to evaluate ontogenetic variations of growth performance, lipid dynamics and nutritional condition of plaice larvae caught in the field during winter 2017 in the eastern English Channel and the Southern Bight of the North Sea. RNA:DNA ratios showed that larvae at the beginning of metamorphosis (stage 4) had a lower growth rate than younger individuals (stages 2 and 3). A significant increase in the proportion of triglycerides also occurred at stage 4, indicating energy storage. Histological indices indicated that most of the larvae were in good condition, even younger ones with low lipid reserves. There was, however, an increase in the proportion of healthy individuals over ontogeny, especially with respect to liver vacuoles which were larger and more numerous for stage 4 larvae. Combined together, these condition indices revealed the ontogenetic shift in the energy allocation strategy of plaice larvae. Young larvae (stages 2 and 3) primarily allocate energy towards somatic growth. The decrease in growth performance for stage 4 was not related to poor condition, but linked to a higher proportion of energy stored as lipids. Since the quantity of lipid reserves is particularly important for plaice larvae to withstand starvation during metamorphosis, this could be considered as a second critical period after the one of exogenous feeding for larval survival and recruitment success.

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