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Map showing the Connecticut River drainage area including the four river sampling sites (stars) for American shad located at km 1 (Old Lyme, Conn.), km 139 (Holyoke, Mass.), km 198 (Turners Falls, Mass.), and km 228 (Vernon, Vt.).
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We examined total and tissue-specific energy content of upstream-migrating American shad (Alosa sapidissima) in the Connecticut River. Total energy depletion over the course of the 228-km migration ranged from 35 to 60%. The approximate contributions of different tissues to energy use during migration were white muscle 57%, subdermal fat 27%, red m...
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... energy storage areas. We also examine the impact of migration distance, fish size, and sex on total and tissue-specific energy use. This allows us to assess the overall cost of migration in American shad and to gain an understanding of how the increased metabolic costs of migration are apportioned between different tissues and energy supplies. (Fig. 1). Efforts were made to sample fish within 1 week of the arrival of the first migratory cohort at each location to minimize the confounding effects of variable amounts of time spent in the river prior to sampling. It is difficult to know precisely when fish first enter the estuary, and fish ladders are not necessar- ily operated with ...
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Citations
... With northern populations displaying lower spawning mortality rates and a greater instance of repeat spawning than southern populations, nutrient delivery through excretion and spawning (gametes) may be most important. American shad generally do not feed upon entering rivers, but their mass loss during the run can be considerable (Glebe and Leggett, 1981a,b;Leonard and McCormick, 1999;Walter and Olney, 2003). Regardless of the source, the level to which the historical delivery of marine nutrients, and the seaward return of nutrients by juveniles, has shifted because of dams. ...
... We applied average values of 36 % and 46% mass loss for males ( M ) and females ( F ) respectively ( Table 2). We acknowledge this as a simplifying assumption as it is well-known that distance traveled (Leonard and McCormick, 1999), temperature experience (Glebe and Leggett, 1981b;Raabe and Hightower, 2014) and residence time (Raabe and Hightower, 2014) all influence the extent of individual mass lost in freshwater. ...
... For the gonads, however, we note that the testes are enriched in both N and P (∼10 % over other tissues) while the ovaries are enriched in N (∼10%) but depleted in P (∼50%), resulting in an over estimate for P. However, because female gonadal mass represents approximately 10 % of the mass of a prespawn female (Leonard and McCormick, 1999), and the applied mass loss based on observations is more than four-fold greater (46% mass loss during spawning), the influence on phosphorus estimates is minimal. ...
American shad (Alosa sapidissima) are native to the east coast of North America from the St. Johns River, Florida, to the St. Lawrence River region in Canada. Since the 1800s, dams have reduced access to spawning habitat. To assess the impact of dams, we estimated the historically accessed spawning habitat in coastal rivers (485,618 river segments with 21,113 current dams) based on (i) width, (ii) distance from seawater, and (iii) slope (to exclude natural barriers to migration) combined with local knowledge. Estimated habitat available prior to dam construction (2,752 km²) was 41% greater than current fully accessible habitat (1,639 km²). River-specific population models were developed using habitat estimates and latitudinally appropriate life history parameters (e.g., size at age, maturity, iteroparity). Estimated coast-wide annual production potential was 69.1 million spawners compared with a dammed scenario (41.8 million spawners). Even with optimistic fish passage performance assumed for all dams (even if passage is completely absent), the dam-imposed deficit was alleviated by fewer than 3 million spawners. We estimate that in rivers modeled without dams, 98,000 metric tons of marine sourced biomass and nutrients were annually delivered, 60% of which was retained through carcasses, gametes and metabolic waste. Damming is estimated to have reduced this by more than one third. Based on our results, dams represent a significant and acute constraint to the population and, with other human impacts, reduce the fishery potential and ecological services attributed to the species.
... So, efforts should be invested to deal with the spatiotemporal complexity of migration and go beyond univariate analysis by integrating multiple cues simultaneously. If implemented, this improvement would also require venturing into the field of energetic expenditures associated with migration and the subsequent existing trade-offs between investments in migration or reproduction (Leonard and McCormick 1999;Morbey and Hedenström 2020). In addition, no additional mortality was considered during migration despite the fact that these rivers are highly anthropized. ...
We developed the HoOS simulator (Hasty or Omniscient Shads) to assess how upstream migration and reproductive timings influence allis shad Alosa alosa (Linnaeus, 1758) temperature-dependent survival of early-life stages. We tested the influence of two reproductive timings depending on whether spawners can synchronize their spawning activity (i.e., referred to as “omniscient fish”) with optimal temperature for early-life stage survival, or simply spawn immediately after arriving on the spawning grounds (i.e., referred to as “hasty fish”). We computed the survival of early-life stages using an existing relationship that predicts the survival of allis shad from hatch to up to 14 days post-hatching according to the time series of daily water temperature at the spawning grounds. The early-stage survival rate was then calculated for each reproductive timing using a range of upstream migration speed while also considering physiological constraints to spawning adults on reproductive timing. Results indicated that the temperature-dependent survival of early-life stages relative to the migration timing by the adults was dome-shaped, confirming the hypothesis of an optimal period for reproduction across the reproductive season. The expected gain of the early-stage survival under the “omniscient” reproductive timing was highest for early arrivals of adults on the spawning grounds, lowest for intermediate arrivals, and was increasing for late arrivals. The HoOS simulator demonstrated that early and late arrivals of adults calculated as less favorable could be compensated by an “omniscient” reproductive timing. In contrast, for intermediate transit times, the two spawning timings performed the same. Our results provided modelling support that a good match of spawning activity with optimal thermal conditions is particularly crucial for the early arriving adults to maximize allis shad fitness-related outcomes (here early-stage temperature dependent survival), a causal relationship frequently claimed but rarely explored.
... Various factors that can influence migratory timing have been identified in previous work [78,91]. Among these are the energy reserves required for migration, which may be especially important in terrestrial movement and if migratory costs are dependent on an interaction between size and time [92]. However, the migration distances in our study system (stream to lake) are generally in the range of meters to no more than a few kilometers, and so both downstream and upstream migration can be achieved with minimal energy investment. ...
Background:
Seasonal spatio-temporal variation in habitat quality and abiotic conditions leads to animals migrating between different environments around the world. Whereas mean population timing of migration is often fairly well understood, explanations for variation in migratory timing within populations are often lacking. Condition-dependent tradeoffs may be an understudied mechanism that can explain this differential migration. While fixed condition-specific thresholds have been identified in earlier work on ontogenetic niche shifts, they are rare in differential migration, suggesting that thresholds in such systems can shift based on temporally variable environmental conditions.
Methods:
We introduced a model based on size-specific tradeoffs between migration and growth in seasonal environments. We focused on optimal migratory timing for first-time migrants with no knowledge of an alternative habitat, which is a crucial stage in the life history of migratory salmonids. We predicted that optimal timing would occur when individuals move from their natal habitats based on a seasonally variable ratio of predation and growth. When the ratio becomes slightly more favorable in the alternative habitat, migratory movement can occur. As it keeps shifting throughout the season, the threshold for migration is variable, allowing smaller individuals to move at later dates. We compared our model predictions to empirical data on 3 years of migratory movement of more than 800 juvenile trout of varying size from natal to feeding habitat.
Results:
Both our model and empirical data showed that large individuals, which are assumed to have a lower predation risk in the migratory habitat, move earlier in the season than smaller individuals, whose predicted predation-to-growth ratio shifted to being favorable only later in the migratory season. Our model also predicted that the observed difference in migratory timing between large and small migrants occurred most often at low values of growth differential between the two habitats, suggesting that it was not merely high growth potential but rather the tradeoff between predation and growth that shaped differential migration patterns.
Conclusions:
We showed the importance of considering condition-specific tradeoffs for understanding temporal population dynamics in spatially structured landscapes. Rather than assuming a fixed threshold, which appears to be absent based on previous work on salmonids, we showed that the body-size threshold for migration changed temporally throughout the season. This allowed increasingly smaller individuals to migrate when growth conditions peaked in the migratory habitat. Our model illuminates an understudied aspect of predation as part of a condition-dependent tradeoff that shapes migratory patterns, and our empirical data back patterns predicted by this model.
... These individuals and their progeny then move downstream to the estuary after spawning (Zhang, 2001). The whole process of long-distance migration and spawning is energetically costly, particularly in adverse river environmental conditions (Bernatchez & Dodson, 1987;Leonard & McCormick, 1999). In spite of the costs, grenadier anchovy does not usually ingest food during migration (Yuan, 1978) and must withstand prolonged periods of food deprivation. ...
In this study, we investigated the activity levels of two major digestive enzymes (pepsin and lipase) in the commercially important Japanese grenadier anchovy Coilia nasus during its upstream migration to analyse the digestive physiological responses to starvation and to analyse the influence of the water temperature on enzyme activity. Water temperature had a significant effect on pepsin activity, while long‐term starvation resulted in a significant decrease in pepsin activity. As starvation continued, however, a slight increase in pepsin activity between the Wuhu (440 river km) and Anqing (620 river km) regions may indicate that C. nasus had refeeding behaviour due to its large expenditure of energy reserves. In contrast, lipase activity was not significantly affected by the water temperature but the effect of fasting increased as much as 13% of lipase activity from the Chongming region (20 river km) to Anqing region, suggesting that the stored lipids of grenadier anchovy were mobilised to meet energy requirements of upstream migration activity and gonad development. Lipid mobilisation activated lipoprotein lipase (LPL; proteins with lipase activity) to hydrolyse triacylglycerides (TAG), which is the first step of lipid assimilation and obtained energy from fatty acids under fasting conditions. Therefore, the increased lipase activity is attributed mainly to the lipase that is involved in endogenous lipid hydrolysis. Grenadier anchovy appears to adapt to long‐term starvation during migration and the increased lipase activity may indicate a crucial effect on lipid metabolism. This study demonstrated that distinct alterations occur in pepsin and lipase activities during the spawning migration of grenadier anchovy due to exogenous nutrition and endogenous metabolism. Furthermore, it provides a basis for further research on the digestive physiology and energy metabolism in this species.
... It is known that migratory fishes that spawn once in a life time often imply a great energy investment prior to spawning and rely entirely on endogenous energy stores to fuel return migration 28 . In accordance with the observations in American shad (Alosa sapidissim) 29 and chum salmon 30 , the decreases of lipid and protein contents and increase of glycogenic amino acid in yellowfin goby muscle reflect a physiological role of gluconeogenesis occurred in muscle in feeding the gonad during the long distance migration. ...
Data-driven approaches were applied to investigate the temporal and spatial changes of 1,022 individuals of wild yellowfin goby and its potential interaction with the estuarine environment in Japan. Nuclear magnetic resonance (NMR)-based metabolomics revealed that growth stage is a primary factor affecting muscle metabolism. Then, the metabolic, elemental and microbial profiles of the pooled samples generated according to either the same habitat or sampling season as well as the river water and sediment samples from their habitats were measured using NMR spectra, inductively coupled plasma optical emission spectrometry and next-generation 16 S rRNA gene sequencing. Hidden interactions in the integrated datasets such as the potential role of intestinal bacteria in the control of spawning migration, essential amino acids and fatty acids synthesis in wild yellowfin goby were further extracted using correlation clustering and market basket analysis-generated networks. Importantly, our systematic analysis of both the seasonal and latitudinal variations in metabolome, ionome and microbiome of wild yellowfin goby pointed out that the environmental factors such as the temperature play important roles in regulating the body homeostasis of wild fish.
... The condition and sex-specific differences in hickory shad on spawning grounds in the Susquehanna River and elsewhere are the result of a culmination of stresses and physiological changes that occur in fish in preparation of spawning activity during a long distance migration from the Atlantic Ocean. Upstream migration is energetically demanding and stressful, perhaps more so for female shad (Leonard & McCormick 1999a, Leonard & McCormick 1999b. ...
Background:
Declines in Hickory shad (Alosa mediocris) populations in Chesapeake Bay have prompted efforts at captive propagation of wild broodfish for stock enhancement and research.
Objective:
The objectives of this study were to evaluate injuries sustained, and immediate and delayed (24 hours) effects on blood variables related to 2 fish capturing methods (electrofishing [EF] and angling).
Methods:
Blood specimens were collected from fish immediately following capture by EF and angling (n = 40 per sex and capture method) from the Susquehanna River (MD, USA). Additional fish (n = 25 per sex and capture method) were collected on the same day, placed in holding tanks and bled 24 hours following capture. Blood data that were non-Gaussian in distribution were transformed (Box-Cox), and effects of sex, method of capture, and holding time were tested using ANOVA with general linear models. Fish were evaluated for injuries by necropsy and radiography.
Results:
Sex-specific differences were observed for RBC, HGB, PCV, MCH, MCHC, total proteins (TP), globulins, glucose, calcium, AST, CK, and lactate, while RBC, HGB, PCV, MCV, MCH, MCHC, TP, albumin, globulins, glucose, potassium, sodium, AST, CK, and lactate differed significantly by fish capturing method. Electrofishing may have induced greater disruption in blood variables, but mortality (4%) was not significantly different compared to angling.
Conclusions:
Electrofishing for Hickory shad using a constant DC voltage resulted in numerous hematologic and biochemical changes, with no additional injuries or deaths compared to angling. Capture method must be considered when evaluating fish condition, and blood variables should be partitioned by sex during spawning season.
... The condition and sex-specific differences in hickory shad on spawning grounds in the Susquehanna River and elsewhere are the result of a culmination of stresses and physiological changes that occur in fish in preparation of spawning activity during a long distance migration from the Atlantic Ocean. Upstream migration is energetically demanding and stressful, perhaps more so for female shad (Leonard & McCormick 1999a, Leonard & McCormick 1999b. ...
Anesthetizing fish facilitates handling procedures such as hormone injections to induce spawning, reduces risk of injuries and may reduce stress response. In this study, two experiments were conducted with wild hickory shad Alosa mediocris caught by angling during spawning season; induction and recovery times of fish anesthetized with tricaine methanesulfonate (MS-222, 50–100 mg/L), 2-Phenoxyethanol (PhE, 25–400 mg/L) and eugenol (Aqui-s, 25–100 mg/L) were compared in the first experiment, while physiological responses in fish following anesthesia were compared in the second experiment. Total time to induce and recover from anesthesia in fish was lowest with 100 mg/L MS-222 and 100 mg/L eugenol. External signs of stress (muscle spasms, coughing, decreased respiration) were evident in fish during anesthesia with highest concentration of PhE (400 mg/L), while physiological indicators of toxicity were evident in fish anesthetized with a lower concentration of PhE (100 mg/L) compared to controls (decreased total proteins and increased hemolysis, plasma lactate and glucose). Anesthesia with MS-222 (100 mg/L) and eugenol (100 mg/L) resulted in reduced stress response (lower glucose and plasma lactate) 1 h and 6 h post induction. MS-222 and eugenol were effective in rapidly anesthetizing wild hickory shad with fewer signs of stress; however more information is needed regarding effects of these anesthetics on spawning, gamete quality and fertilization success in hickory shad and other species.
Received 16 Dec 2016 accepted 14 Apr 2017 revised 05 Apr 2017
... The fatmeter uses lowpower microwaves to measure subdermal lipid content in a non-destructive manner (Kent 1990, Crossin & Hinch 2005. In some fish species, subdermal lipid reserves are preferentially used to support migration (Leonard & McCormick 1999). ...
... The increase in subdermal lipid stores observed prior to egress, particularly for juvenile Atlantic croaker, is consistent with the use of subdermal lipids to support migration (as seen in American shad; Leonard & McCormick 1999). If subdermal lipids are also used by juvenile summer flounder to undertake coastal migrations in the late fall, then a portion of the juveniles from Chesapeake Bay may not have sufficient energy to complete these directed movements (e.g. the large proportion of low-lipid individuals observed in the 2012 year class; Fig. 3). ...
The condition of individuals in a year class may contribute to recruitment variability due to differential survival of poor- and well-conditioned fish, but the temporal dynamics of juvenile fish condition are poorly understood. We examined inter- and intra-annual dynamics of condition for juveniles of 3 species collected from estuarine nursery areas of Chesapeake Bay from November 2010 to June 2014. We describe temporal patterns in length-based indices, the hepatosomatic index (HSI), and relative subdermal lipid estimates for juvenile summer flounder Paralichthys dentatus (n = 1771), Atlantic croaker Micropogonias undulatus (n = 3911), and striped bass Morone saxatilis (n = 874). Multiple indices provided a more complete understanding of energy-storage strategies for juveniles because temporal patterns among condition indices were not congruent for a given species. Most juvenile summer flounder and Atlantic croaker migrate from Chesapeake Bay in the fall, and both species exhibited increases in subdermal lipids in the time period prior to migration. For all species, individuals that remained in the estuary during winter exhibited high HSI values, indicating a common energy-storage strategy during winter. Mean condition of juveniles varied among year classes, but differences were inconsistent among indices, suggesting that energy was differentially stored among tissues for these year classes. Densitydependent effects contributed to variation in mean condition for summer flounder and striped bass. Our understanding of recruitment variability may be improved by assessing annual differences in mean condition as revealed by multiple indices.
... The occurrence of AJ at S1 in contrast to the absence of AH plus the spatial segregation of young and adult specimens of AJ in contrast to the absence of such pattern among the AH population, indicate the ability of the latter in transposing barriers and corroborates the proposition that, in opposition to AH, AJ is a migratory species (Mazzoni et al., 2004). Thus, assuming that reproduction is a driving force for the ascent of adult fishes and that ascending involves a high loss of somatic energy (Dodson, 1997;Leonard and McCormick, 1999) AJ spends more energy than AH, when such aspect of reproductive strategy (migration) is considered. In fact, it was estimated that the costs with OI (egg size and egg numbers) are equal for both species but the overall strategy, the above parameter plus breeding time, is 25% more expensive in AH. ...
The reproductive strategy of two congeneric species (Astyanax janeiroensis--AJ and Astyanax hastatus--AH) was assessed, in order to address the costs imposed by different investment patterns, on four major life history characters: migratory behaviour, breeding time, fecundity and egg size. Altitudinal segregation of young and adult individuals plus data from the literature indicated that AJ is a migratory species, whereas AH is non migratory. Breeding season (BS) analysis revealed that AH was reproductively active year-round (12 months), whereas AJ breeds during 9 months. Brood costs (Ovarian Investment) were almost the same for both species (AJ--364 oocytes/g and egg volume = 0.52 mm(3); AH--702 oocytes/g and egg volume = 0.27 mm(3)). Reproductive costs (RC), considered as product of gamete production (OI) and gamete output (BS), were 1,703.7 for AJ and 2,274.0 for AH; thus RC is 25% higher in AH. It is concluded that the costs with OI (egg size and egg numbers) are equal for both species but as breeding season is larger for AH the overall strategy is 25% more expensive in AH. Thus, it is hypothesized that this 25% should be considered as the extra costs that AJ uses during migration and is compensated by its shorter breeding time (9 months versus 12 months of AH).
More than a century of impoundments in the Penobscot River, Maine, USA, has contributed to population declines in migratory fish in the system. A decade of change, research, and monitoring has revealed direct and indirect ways that dams have influenced the river habitat, connectivity for migratory fish, and the food web. The removal of two main-stem dams (in 2012 and 2013) and bolstering of fish passage have been part of coordinated restoration efforts in the watershed. Integral to this undertaking was support for short- and long-term monitoring and research that included physical habitat, fish passage, and broad scale ecological assessments. Herein we discuss the seven interconnected and complex ways that dams have affected the Penobscot River ecosystem, particularly for migratory fish. These include familiar influences ascribed to dams: i) impaired access to habitat, ii) injury and mortality, and iii) delays of migration. Other ecological influences are less studied and more subtle: iv) facilitation of predation, v) community shifts, and vi) demographic shifts. Lastly, dams result in vii) a loss of ecosystem services that would otherwise be intact in an unimpounded system. We draw on both direct examples from the Penobscot River and broader information to characterize how impoundments have transformed this ecosystem for more than a century. Recent dam removals and mitigation efforts have reestablished some of these ecological functions.
