Alternative mating strategies in Atlantic salmon and brown trout

By screening variable number of tandem repeat (VNTR) loci, multiple paternity within clutches has been found in wild populations of southern European Atlantic salmon (Salmo salar) and brown trout (Salmo trutta). For Atlantic salmon, we determined the relative contribution of alternative male phenotypes to the next generation. Individual males that are morphologically juvenile yet sexually mature fertilized a large proportion of eggs, and they thereby contributed to an increase of genetic variability in wild populations via (1) balancing the sex ratio, (2) increasing outbreeding, and (3) enlarging the effective population size, in part a consequence of (1) and (2). In addition, these precocious males ensured that interspecific spawns involving Atlantic salmon females and brown trout males (a fairly common occurrence in southern Europe where the two species are sympatric) resulted mostly in Atlantic salmon progeny. For brown trout, preliminary genetic results indicated that multiple paternity, when present, was not due to alternative mating strategies by males, but rather to successive fertilizations by adult suitors.     

Sexual Selection and Breeding Patterns: Insights from Salmonids (Salmonidae)

Although "intrasexual selection" has been accepted as the mechanism by which males evolve elaborate secondary sexual traits which are used in aggressive contests, the importance of "intersexual selection" as a mechanism by which males have acquired exaggerated traits to display to females during courtship was less readily accepted. In spite of this scepticism, several genetic models have supported the latter idea, and many empirical studies showed that females were generally more discriminating in mate choice than males, because of differences in relative investment between sexes. Nowadays, this idea is reinforced by various concepts (parental investment, potential reproductive rate, environmental potential for polygamy...) which stress that the strength of sexual selection is related to many interdependent factors, such as mating systems, resource distribution (food, habitat, mate), life history and other ecological characteristics. The case of Salmonids is presented here to show how novel information on sexual selection has contributed to the understanding of the plasticity of breeding patterns in the context of evolutionary biology.     

Interspecific barriers between salmonids when hybridisation is due to sneak mating

Male sneaking behaviour can lead to interspecific hybridisation if sneakers attempt to fertilise ova in heterospecific mating, contributing to break down of interspecific barriers. In south European rivers, sneaking Atlantic salmon males fertilise an important proportion of ova from adult females in heterospecific crosses, up to 65%. In a south French flow-controlled stream, we found that they were able to naturally fertilise brown trout ova in absence of brown trout males. Aggressiveness of brown trout males towards sneaking salmon males and low survival of hybrids issued from salmon sneakers are found to be interspecific barriers.     

Variation of reproductive behaviour and success of males adopting different tactics in Atlantic salmon

  The use of video photography normally involves sub‐aqua recording via a camcorder device. The problem is that the scientist needs to be directly involved in the recording underwater or else viewing the results of the footage after the event. The system we have developed enables real time images to be relayed to a boat, satellite of shore‐based recorder. With a two‐way audio communication system, it is possible for the scientist/team to direct the camera work of the divers, remotely. The configuration of this system are discussed, and a wide range of scientific applications suggested.     

Nest Placement and Egg Distribution in Atlantic Salmon Redds

Atlantic salmon, Salmo salar, deposit their eggs in excavated depressions called nests. These nests are built from downstream to upstream within one or more redds, and each redd corresponds to a continuous area of the streambed disturbed by the female digging activities. Redd topographic measurements and egg excavation were performed to determine number of nests per redd and per female, nest depth, distances between successive nests, number of eggs deposited per nest, and egg survival in nests created by six grilse Atlantic salmon, five spawning in an experimental stream (Lapitxuri channel) and one in a natural stream (Lurgorrieta Creek, a tributary of the Nivelle River in southwest France). All females constructed a single redd, except one which built two redds in the channel. Redd surface area ranged between 2.3 and 5.7m². Each redd had a raised mound of gravel or dome under which most of the eggs were located, and an upstream depression or pot. Based on expected egg-to-juvenile survival rates previously obtained in the Lapitxuri channel and on juvenile recoveries, between 96 and 97% of the eggs deposited in the channel sections were retrieved. Each female constructed 7 to 11 nests over a period of 3 to 5 days. The first three nests had an average burial depth of 12.9cm (±1.6SD) which was greater than the last three nests (mean 9.5cm±2.6SD). Eggs removed from the first three nests had higher fertility rates (95.5% vs. 87.2%), greater survival (83.5% vs. 63.1%) and lower occurrences of abnormalities (1.9% vs. 5.5%) than those deposited in the last three nests. Typically, the percentage of eggs deposited per female decreased from the first to the last nest, such that the last two to three nests possessed only a small number of scattered eggs. Similar results were observed in the redd located in Lurgorrieta Creek. The adaptive consequences of the topographic features of redds and the egg allocation patterns we found are discussed.     

Effects of overripening on spawning behaviour and reproductive success of Atlantic salmon females spawning in a controlled flow channel

The influence of overripening on salmonid egg quality has often been described in artificial breeding. However, no information exists on the impact of this factor on fish reproducing naturally, although it is likely to be critical, especially in disturbed environments. Pairs of Atlantic salmon in which the size of the male and the female was evenly matched were released in a spawning channel from 1 to 12 days after ovulation. Handling did not affect egg quality. Overripening had a negative effect on egg viability. Egg mortality, infertility and malformations increased with the number of days after ovulation. In females which had just matured, 16·7% of their recovered eggs were dead, 3·1% were unfertilized and 1·7% were malformed, as compared to 25·4, 9·7 and 5·4%, respectively, in females that had been mature for at least a week. Overripening played a role in both female spawning behaviour and capacity to spawn, as females which had been mature for over a week released their eggs more quickly while they retained more ova in their abdominal cavity than early spawning females. Thus, in studies of reproductive behaviour and in particular of sexual selection, it is critical to take into account the maturation status of females since the duration of spawning activity is used by various authors to determine female sexual motivation. Alternatively, overripening may decrease the female selectivity for mates.     

Multiple paternity increases effective size of southern Atlantic salmon populations

Genetic analyses were performed on the progeny of Atlantic salmon (Salmo salar L.) sampled in natural redds of three rivers flowing into the Bay of Biscay, the Nivelle, the Mandeo and the Sella. These rivers are at the southern limit of the European distribution of the species and their populations are small and endangered by human activities. Nine variable number of tandem repeat (VNTR) loci (five minisatellites and four microsatellites) were used for parentage analysis. Multiple male participation was recognized in the fertilization of eggs. A large proportion was fertilized by precociously mature parr. We demonstrate that multiple paternity derived from mature parr is crucial for the conservation of genetic variability in small populations of Atlantic salmon.     

Effects of courtship and relative mate size on sexual motivation in Atlantic salmon

Both sexes of Atlantic salmon Salmo salar were very sensitive to the absence of their mate on the spawning ground, particularly females during the last hour before oviposition. The improvement of behavioural chaining just before viposition might elicit the accurate timing of synchronized gamete release for successful fertilization. While the reproductive behaviour of the opposite sex could largely affect breeding activity, relative mate size appeared to be the prevailing sexual motivation factor in this species. Even in the absence of courtship, large mate size may constitute a supra-stimulus inducing an increase in spawning behaviour of the other sex. Females with smaller males delayed their first spawning activity, took longer to spawn and made more nests than those with large males. However, female egg retention was not influenced by relative male size.