Substrate selection by juvenile Atlantic cod (Gadus morhua): effects of predation risk

Although predator avoidance has been proposed as one possible factor influencing the distribution of fish among substrate types, no study has addressed this question directly. Groups of juvenile Atlantic cod were offered a choice between pairs of the following three substrates: sand, gravel-pebble and cobble. Their distribution on these substrates was compared prior to, during and following exposure to a predator (i.e. a larger conspecific). With no apparent risk of predation, juvenile cod preferred sand or gravel-pebble. When cobble was present, juveniles hid in the interstitial spaces of this substrate in the presence of a predator. With no cobble present, juveniles showed no preference between sand and gravel-pebble, and did not seek refuge from predation in association with these substrates. Following exposure to a predator (i.e. 2.5 h later) larger juvenile cod again showed a preference for the finer-grained substrates, but smaller individuals continued to associated with the cobble. The presence of cobble resulted in fewer juveniles being captured and a significant increase in the latency until the first juvenile was captured by the predator. Results are discussed with respect to the effects of predation on the distribution and survival of fishes among substrate types.     

Influence of cruising and ambush predators on 3-dimensional habitat use in age 0 juvenile Atlantic cod Gadus morhua

Predators and prey often co-exist at high densities within the same habitat, yet the behavioural and spatial dynamics underlying this co-existence are not well known. To better understand small-scale, predator-prey co-occurrence, the spatial patterns and behaviour of age 0 juvenile cod Gadus morhua 75-88 mm SL and two of their known predators, age 2+ cod and short-horn sculpin Myoxocephalus scorpinus, were examined in two habitats (i.e., sand and eelgrass) using three-dimensional video analysis. Both habitat and predator type interacted to result in unique spatial patterns of prey. Spatial overlap between predators and prey was highest in open habitat in the presence of the cruising predator but lowest in the presence of sculpin in the same habitat. In eelgrass, age 0 cod avoided predators primarily along the vertical axis (i.e., distance off bottom). Age 0 cod stayed above eelgrass in the presence of sculpin but lowered themselves into the eelgrass while in the presence of predator cod. Anti-predator behaviour (i.e., predator-prey distance, prey cohesion and freezing) was significantly reduced over eelgrass compared to sand, suggesting eelgrass has lower ‘inherent risk’ than open habitats. However, predator consumption was similar across all treatments, suggesting that, 1) complex habitat also impairs the visual cues needed for anti-predator behaviour (e.g., schooling) and assessing the location of predators, and 2) predators change their behaviour with habitat to enhance their opportunities for finding and capturing prey.     

Shallow water predation risk for a juvenile flatfish (winter flounder; Pseudopleuronectes americanus, Walbaum) in a northwest Atlantic estuary

Many small fish, including several juvenile Atlantic flatfish, are most abundant in shallow areas presumable because these habitats enhance survivorship and/or growth. In this study, we investigated size-dependent depth distributions and the role of shallow habitats as predator refuges for age-0 winter flounder (Pseudopleuronectes americanus) in a northwest Atlantic estuarine nursery. Analysis of trawl surveys performed during the larval settlement period throughout the Navesink River and Sandy Hook Bay, New Jersey, showed that as fish increased in size, depth of occurrence gradually decreased, so that individuals >35 mm standard length (SL) were concentrated in habitats ∼1 m deep. Tethering in structurally simple and adjacent shallow and deep habitats showed that predation risk for flounder (30-50 mm SL) was low in shallow water (<1 m) and increased rapidly with depth. Summer flounder (Paralychthys dentatus), which were more abundant in trammel nets in deep habitats and included winter flounder in their diets, appeared to be important consumers of tethered fish. Our results indicate that following larval settlement, winter flounder emigrate from or suffer high mortality in deeper water to become concentrated in shallow habitats that can serve as predator refuges even when they lack complex physical structures. These results highlight the potential for functional habitat loss when natural and/or anthropogenic factors make shallow habitats unavailable to young fish.     

The effect of growth rate,size, and season on oocyte development and maturity of Atlantic cod (Gadus morhua L.)

The effect of growth rate, body weight, age, and season on ovarian development and maturation was investigated for Atlantic cod (Gadus morhua L.) reared in the laboratory over 10 months, for each of two consecutive years, 1978–1980. Cod were also collected from the Gulf of St. Lawrence, the Scotian Shelf, Georges Bank, the Flemish Cap, and the N.E. Newfoundland Shelf.The state of maturity was recognized by oocyte size. Stage I oocytes did not vary in size with growth rate, season, age, or maturity, while the size of stage II oocytes was positively correlated with maturity and negatively correlated with maximum stage of development achieved. Ovarian wall thickness was positively correlated with age and maturity.The frequency distribution of stage I and II oocytes distinguished the state of maturation, with cod that would mature by the next spawning season having a minimum of 20% ($\text{x}\text{?}\text{= 42\%}$) of their oocytes at stage II or greater level of development.Maturing 3-yr-old cod had greater life specific growth rates than immature 3-yr-olds, but growth rates during the third year itself were not significantly different. A hypothesis of a three-part density-dependent mechanism controlling fecundity is postulated. Future reductions in partial recruitment and total fecundity are predicted for the Gulf of St. Lawrence cod stock based on calculated growth rates for Gulf cod in 1979.     

Use of space and habitats by meadow voles at the home range,patch and landscape scales

Using capture/recapture methods, we examined the spatial usage patterns of Microtus pennsylvanicus within and between experimentally created habitat patches of three sizes (1.0, 0.25 and 0.0625 ha) and between a 20-ha fragmented and a 20-ha continuous habitat landscape. We tested the prediction that home ranges near patch edges would be qualitatively different from those in patch interiors, and that the edge:interior habitat ratio could be used to make predictions concerning the dispersion and spatial use of individuals occupying different sized patches and between landscapes with different habitat structure. We found adult females on patch edges to have larger and more exclusive home ranges, larger body sizes, longer residence times, and to reproduce at a higher frequency than those in patch interiors. These edge effects also appeared to be largely responsible for the greater proportion of larger, reproductive females we found in small than larger patches and in the fragmented than in the continuous habitat (control) landscape. The selection of higher quality edge habitats by dominant females and the relegation of sub-dominants to patch interiors provides an explanation for the observed differences in the distribution and performance of females over patches and between landscapes.     

捕食风险对越冬根田鼠肠道寄生物易感性的影响

在自然生态系统中,不同营养级物种可通过特征介导间接效应对生态系统的稳定及种群产生深刻的影响。但目前有关特征介导间接效应的实验研究多见于无脊椎动物、鱼类和两爬类。本研究以根田鼠为对象,在野外围栏内建立预防捕食者和未预防捕食者两种实验处理种群,并通过采用麦克马斯特法测定两种处理种群实验个体肠道寄生物感染种类及感染率和感染强度,采用PHA（phytohemagglutinin）反应和白细胞分类计数测定不同处理种群实验个体免疫能力,以分析捕食风险对根田鼠肠道寄生物的感染效应。结果表明,未预防捕食者处理组根田鼠PHA反应、白细胞计数和淋巴细胞计数较预防捕食者处理组实验个体显著降低,而球虫 E. wenrichi 的感染率和感染强度则显著增加,但绦虫和线虫以及其他3种球虫的感染率和感染强度无显著差异。结果表明,捕食者可通过介导猎物免疫力特征而间接影响猎物肠道寄生物的感染,验证了本项提出的捕食风险可通过降低根田鼠的免疫能力而增加其肠道寄生物感染的假设。     

A SNP/microsatellite genetic linkage map of the Atlantic cod (Gadus morhua)

A first genetic linkage map of the Atlantic cod ( Gadus morhua ) was produced, based on segregation data from 12 full-sib families of Norwegian origin. The map contained 174 single nucleotide polymorphism markers and 33 microsatellites, distributed on 25 linkage groups and had a length of 1225 cM. A significant difference in recombination rates between sexes was found, the average ratio of female:male recombination rates being 1.78 ± 1.62 (SD).     

Nest site preferences of the Woodlark (Lullula arborea) and its association with artificial nest predation

The Woodlark is an insectivorous bird, which is listed as a priority species in Switzerland. In Valais, a stronghold of this species in the country, the birds breed in intensively managed vineyards and show a preference for parcels with ground vegetation during territory establishment. As a ground-breeder, the species is highly vulnerable to nest predation by avian and mammal predators. The aims of our study were firstly to investigate nest site preferences of the woodlark within vineyards and secondly to compare the predation risk of artificial nests dependent of ground vegetation structure. Our results point out that the Woodlark prefers patches of tall and dense ground cover within vegetated vineyard parcels and avoids parcels that have been treated with herbicides. In a follow-up experiment we conducted a study comparing the predation rate of artificial nests between bare parcels (<20% vegetated area) and vegetated parcels (>40% vegetated area). Artificial nests equipped with one quail egg were distributed pairwise between two adjacent parcels that fulfilled the upper criteria and were monitored by trail cameras during 10–12 days. Predation rate was generally low (4 predation events) and only occurred in bare parcels. These data indicate that conspicuousness of avian nests may be decreased in vegetated parcels and that the amount of vegetation can lower the predation risk on ground breeding birds - another indication for the importance of ground vegetation for a successful conservation of the endangered Woodlark in Swiss vineyards.     

Using experimental ecology to understand stock enhancement: Comparisons of habitat-related predation on wild and hatchery-reared Penaeus plebejus Hess

Marine stock enhancement is often characterized by poor survival of hatchery-reared individuals due to deficiencies in their fitness, such as a diminished capacity to avoid predators. Field experiments were used to examine predation on Penaeus plebejus, a current candidate for stock enhancement in Australia. We compared overall survival of, and rates of predation on, wild P. plebejus juveniles, naïve hatchery-reared juveniles (which represented the state of individuals intended for stock enhancement) and experienced hatchery-reared juveniles (which had been exposed to natural predatory stimuli). Predation was examined in the presence of an ambush predator (Centropogon australis White, 1790) and an active-pursuit predator (Metapenaeus macleayi Haswell) within both complex (artificial macrophyte) and simple (bare sand and mud) habitats. Overall survival was lower and rates of predation were higher in simple habitats compared to complex habitats in the presence of C. australis. However, the three categories of juveniles survived at similar proportions and suffered similar rates of predation within each individual habitat. No differences in survival and rates of predation were detected among habitats or the categories of juveniles when M. macleayi was used as a predator. These results indicate that wild and hatchery-reared P. plebejus juveniles are equally capable of avoiding predators. Furthermore, exposure of hatchery-reared juveniles to wild conditions does not increase their ability to avoid predators, suggesting an innate rather than learned anti-predator response. The lower predation by C. australis in complex habitats was attributed to a reduction in this ambush predator's foraging efficiency due to the presence of structure. Ecological experiments comparing wild and hatchery-reared individuals should precede all stock enhancement programs because they may identify deficits in hatchery-reared animals that could be mitigated to optimize survival. Such studies can also identify weaknesses in wild animals, relative to hatchery-reared individuals, that may lead to the loss of resident populations.     

Historical DNA reveals the demographic history of Atlantic cod (Gadus morhua) in medieval and early modern Iceland

Atlantic cod (Gadus morhua) vertebrae from archaeological sites were used to study the history of the Icelandic Atlantic cod population in the time period of 1500–1990. Specifically, we used coalescence modelling to estimate population size and fluctuations from the sequence diversity at the cytochrome b (cytb) and Pantophysin I (PanI) loci. The models are consistent with an expanding population during the warm medieval period, large historical effective population size (N_E), a marked bottleneck event at 1400–1500 and a decrease in N_E in early modern times. The model results are corroborated by the reduction of haplotype and nucleotide variation over time and pairwise population distance as a significant portion of nucleotide variation partitioned across the 1550 time mark. The mean age of the historical fished stock is high in medieval times with a truncation in age in early modern times. The population size crash coincides with a period of known cooling in the North Atlantic, and we conclude that the collapse may be related to climate or climate-induced ecosystem change.     

Bitten down to size: Fish predation determines growth form of the Caribbean coral reef sponge Mycale laevis

Interactions between organisms add complexity to ecosystem function, particularly on coral reefs. The Caribbean orange icing sponge Mycale laevis is semi-cryptic, often growing under coral colonies or between coral branches. This association is reportedly a mutualism, with the sponge deterring boring sponges from invading the coral skeleton and the coral providing an expanding surface for sponge growth. But is there an alternative explanation for the proximity of sponge and coral? We examined the importance of fish predation on the growth of the sponge. While the semi-cryptic growth form of M. laevis predominates on reefs off the Florida Keys and the Bahamas Islands, M. laevis grows with a non-cryptic, erect morphology off Bocas del Toro, Panama. Surveys revealed that sponge-eating fishes were rare or absent at Bocas del Toro compared to sites in the Florida Keys. Because past studies were inconsistent about the palatability of M. laevis to fish predators, we conducted feeding experiments with sponges from all three sites. Crude organic extracts of M. laevis from all three sites were palatable to generalist fish predators in aquarium assays, and field feeding assays and caging experiments conducted in the Florida Keys confirmed that spongivorous fishes readily ate exposed fragments of M. laevis. Our results suggest that M. laevis is restricted to its semi-cryptic growth form by spongivorous predators, with corals providing a physical refuge from predation. This alternative explanation supports the broader hypothesis that Caribbean reef sponges can be categorized on the basis of chemical defense into defended, palatable, and preferred species, the last of which are restricted to refugia.     

Probiotic effect of lactic acid bacteria in the feed on growth and survival of fry of Atlantic cod (Gadus morhua)

A growing concern for the high consumption of antibiotics in aquaculture has initiated a search for alternative methods of disease control. Improved resistance against infectious diseases can be achieved by the use of probiotics. Probiotics are live microorganisms supplemented in food or feed which give beneficial effects on the intestinal microbial balance. In the present study a dry feed containing lactic acid bacteria (Carnobacterium divergens) isolated from Atlantic cod (Gadus morhua)intestines was given to cod fry. After three weeks of feeding the fry was exposed to a virulent strain of Vibrio anguillarum. The death rate was recorded during further three weeks of feeding with lactic acid bacteria supplemented feed. A certain improvement of disease resistance was obtained, and at the end of the experiment lactic acid bacteria dominated the intestinal flora in surviving fish given feed supplemented with lactic acid bacteria. No obvious growth inhibition of V. anguillarum was observed in an in vitro mixed culture of this bacterium and the C. divergens isolated from cod intestines. This revised version was published online in August 2006 with corrections to the Cover Date.     

Characterization of a collagenolytic serine proteinase from the Atlantic cod (gadus morhua)

A collagenolytic proteinase was purified from the intestines of Atlantic cod by (NH₄)₂SO₄ fractionation, hydrophobic interaction chromatography (phenyl-Sepharose) and ion-exchange chromatography (DEAE-Sepharose). The proteinase has an estimated molecular weight of 24.1 (±0.5) kDa as determined by SDS-PAGE and belongs to the chymotrypsin family of serine proteinases. The enzyme cleaves native collagen types I, III, IV and V, and also readily hydrolyzes succinyl-l-Ala-l-Ala-l-Pro-l-Phe-p-nitroanilide (sAAPFpna), an amide substrate of chymotrypsin, as well as succinyl-l-Ala-l-Ala-l-Pro-l-Leu-p-nitroanilide, a reported elastase substrate, but had no detectable activity towards several other substrates of these proteinases or of trypsin. The pH optimum of the enzyme was between pH 8.0 and 9.5 and it was unstable at pH values below 7. Maximal activity of the enzyme when assayed against sAAPFpna was centered between 45 and 50°C. Calcium binding stabilized the cod collagenase against thermal inactivation, but even in the presence of calcium, the enzyme was unstable at temperatures above 30°C.     

Habitat selection of juvenile banana prawns, Penaeus merguiensis de Man: Testing the roles of habitat structure, predators, light phase and prawn size

The effects of fish predators, light phase, habitat structure and prawn size on the habitat preferences of juvenile Penaeus (Fenneropenaeus) merguiensis de Man were examined with laboratory experiments. The behaviour of juvenile P. merguiensis within habitats of different structural complexity was also examined. Experiments were carried out in a tank (1.8 m diameter) divided into four habitats representing: bare substratum, leaf litter (little vertical structure), mangrove pneumatophores (regular vertical structure) and mangrove woody debris (heterogeneous vertical structure). The location of 10 prawns was monitored over 270 min (135 min light and 135 min dark), with different prawns five times for each combination of prawn size class, and predator (no predator; Arius graeffei Kner and Steindachner and Lates calcarifer Bloch).In the absence of predators and during the light phase, when observations on prawn behaviour were made, swimming was the most common behaviour (of seven mutually exclusive behavioural categories) with few differences in behaviour between sizes. All size classes of juvenile P. merguiensis selected vertical structure (mangrove debris and pneumatophores) over low vertical structure (leaf litter and bare substratum), in both light and dark conditions and in the presence or absence of predators. When L. calcarifer was present, the selection by prawns of the mangrove-debris habitat increased significantly. This was attributed to an increase in predation risk in the other habitats. L. calcarifer rarely pursued prey amongst the mangrove-debris structure, compared to habitats with less heterogeneous vertical structure (pneumatophores, leaf litter and bare substratum).     

Temporal and ontogenetic shifts in habitat use of juvenile Pacific cod (Gadus macrocephalus)

Habitat use of age-0 and age-1 juvenile Pacific cod (Gadus macrocephalus) was examined in coastal regions in Kodiak Alaska over daily, seasonal and annual scales. Catch data indicated highly variable recruitment to nursery areas, but a strong separation of distribution by depth among age groups. Age-0 cod were most abundant in the shallows (<3 m) whereas age-1 cod were typically found in depths (9.0-13.5 m). In comparison, age-1 saffron cod (Eleginus gracilis), another highly abundant gadid in the region, were found in shallower depths where age-0 cod often resided. Age-1 cod Pacific cod made diel lateral movements, moving into shallow regions at night where they co-occurred with age-0 cod to a greater extent. Laboratory light-gradient experiments indicated that age-0 cod tolerated intense lighting (~ 20-80 µE m^− 2 s^− 1) typical of shallow water regions whereas larger age-1 Pacific cod strongly avoid bright light given the choice. However, while diet data indicate age-1 cod of both species are moderately piscivorous (3% saffron cod; 16% Pacific cod), we found no direct evidence of predation on smaller conspecific cod, possibly due to the low densities of age-0 cod in the year of the diet study. Together, these data suggest that coastal regions continue to serve a nursery function beyond the 1st year of development for juvenile Pacific gadids, and that small-scale temporal and depth partitioning in these regions is a mechanism by which varying cod species and age classes co-occur.     

A laboratory study of the behavior of two species of grass shrimp (Palaemonetes pugio Holthuis and Palaemonetes vulgaris Holthuis) and the killifish (Fundulus heteroclitus Linneaus)

In this laboratory study, the frequency of locomotor behaviors for two grass shrimp species, Palaemonetes pugio and Palaemonetes vulgaris, and the killifish Fundulus heteroclitus were determined. Additional behaviors, such as grouping of grass shrimp, and pursuit by killifish, the grass shrimp's common predator, were also analyzed. The results show that mean behavioral frequencies for the two shrimp species were rarely significantly different statistically. Each shrimp species altered its behavior differently in the presence of the other shrimp species, but responded similarly to the presence of the predator. The results indicate that the response of each shrimp species to the predator was dependent upon the predator's size and health. In situations involving the large health predator, both grass shrimp species significantly reduced swimming, but did not show a significant difference in walking. Behavioral frequencies of large and small healthy killifish were not significantly different from each other when alone, but were significantly different in the presence of prey, while the large unhealthy killifish's behavior was significantly decreased in all situations compared to both the large and small health killifish.     

Variability in the numbers of post-settlement King George whiting (Sillaginidae: Sillaginodes punctata, Cuvier) in relation to predation, habitat complexity and artificial cage structure

The importance of predation by fish in altering abundances of juvenile King George whiting (Sillaginodes punctata) was examined at multiple locations in Port Phillip Bay, Australia, by manipulating the numbers of piscivorous fish in unvegetated sand and seagrass habitats using cages. Additional information regarding the local abundances of, and habitat use by, the most common piscivorous fish, Western Australian salmon (Arripidae: Arripis truttacea, Cuvier), was gathered using netting surveys and underwater video. Regardless of habitat, abundances of S. punctata were similar in partial cages and uncaged areas. In unvegetated sand, S. punctata were more abundant inside cages than partial cages or uncaged areas. In seagrass, there was no difference in the numbers of S. punctata between caging treatments. Patterns in abundances of S. punctata between cage treatments in each habitat were consistent between sites, but the relative difference in the abundances of S. punctata between habitats was site specific. Abundances of A. truttacea varied significantly between sites, and they consumed a variety of epibenthic fishes including atherinids, clupeids, gobiids, syngnathids and pleuronectids. At one site in Port Phillip Bay (Blairgowrie), A. truttacea occurred more commonly in patches of unvegetated sand than seagrass. Over unvegetated sand, abundances of A. truttacea varied little between partial cages and uncaged areas. The numbers of S. punctata varied between caging treatments and habitats in a manner that was consistent with a model whereby seagrass interferes with foraging by predatory fish and provides juvenile fish with a refuge from predation. The almost total absence of A. truttacea in seagrass habitats and the lack of S. punctata in their diets implies, however, that patterns in S. punctata in seagrass/unvegetated sand mosaics are driven by processes other than direct predation.     

Evidence of a hybrid-zone in Atlantic cod (Gadus morhua) in the Baltic and the Danish Belt Sea revealed by individual admixture analysis

The study of hybrid zones is central to our understanding of the genetic basis of reproductive isolation and speciation, yet very little is known about the extent and significance of hybrid zones in marine fishes. We examined the population structure of cod in the transition area between the North Sea and the Baltic Sea employing nine microsatellite loci. Genetic differentiation between the North Sea sample and the rest increased along a transect to the Baltic proper, with a large increase in level of differentiation occurring in the Western Baltic area. Our objective was to determine whether this pattern was caused purely by varying degrees of mechanical mixing of North Sea and Baltic Sea cod or by interbreeding and formation of a hybrid swarm. Simulation studies revealed that traditional Hardy-Weinberg analysis did not have sufficient power for detection of a Wahlund effect. However, using a model-based clustering method for individual admixture analysis, we were able to demonstrate the existence of intermediate genotypes in all samples from the transition area. Accordingly, our data were explained best by a model of a hybrid swarm flanked by pure nonadmixed populations in the North Sea and the Baltic Sea proper. Significant correlation of gene identities across loci (gametic phase disequilibrium) was found only in a sample from the Western Baltic, suggesting this area as the centre of the apparent hybrid zone. A hybrid zone for cod in the ecotone between the high-saline North Sea and the low-saline Baltic Sea is discussed in relation to its possible origin and maintenance, and in relation to a classical study of haemoglobin variation in cod from the Baltic Sea/Danish Belt Sea, suggesting mixing of two divergent populations without interbreeding.     

Pantophysin (Pan I) locus divergence between inshore v. offshore and northern v. southern populations of Atlantic cod in the north‐east Atlantic

More than 6000 cod Gadus morhua , sampled in coastal and offshore waters stretching from the Barents Sea down to the North Sea, were analysed for frequencies of alleles at the scnDNA pantophysin locus ( Pan I)[formerly called synaptophysin ( Syp I)]. The significant allele frequency difference between the two major stocks of cod in Norway, north‐east Arctic cod (NEAC) and Norwegian coastal cod (NCC), was upheld in all years of the investigation (1993 to 2001), and applied both to larval cod and post‐juveniles of various ages. On a north‐south axis, the appearance of a latitudinal cline of post‐juvenile (≥1 year) allele frequencies was exposed. The intermediate allele frequencies in coastal areas of northern Norway, seem to a large extent to be caused by intermingling of the two stocks, although the existence of populations of coastal cod with alternative Pan I frequencies could not be ruled out. The role of selection is yet unresolved. Depth of the sampling location seemed to have an effect on the allele frequencies and their temporal stability, while there was no indication of seasonal variation in the frequencies. Breeding structure was the most likely cause for upholding the extreme divergence in Pan I frequencies between NEAC and NCC.     

Effect of mother plant age on germination and size of seeds and seedlings in the perennial sedge Carex secalina (Cyperaceae)

The performance of seeds and seedlings in relation to the age of the mother plant was studied in Carex secalina. Seeds of this sedge can differ substantially in size. We planted 100 C. secalina individuals from three populations in a common garden and followed them for four years. We found that mean seed mass varied with plant age, but the pattern of variation was population-specific, with only one population showing significant reduction in seed mass with age. Similarly, germination frequency changed with age differently in different populations. The relationship between the age of the mother plant and the height of emerged seedlings did not differ between populations. In spite of the fact that plant size and mean seed mass exhibited similar patterns of variation within populations, there was no correlation between these two variables at the level of individuals. This means that the size of C. secalina tufts does not determine how big the seeds will be. Moreover, there was no relationship between mean seed mass and the height of seedlings. Presumably, factors intrinsic to each plant determine the production of either small or large seeds in a population-specific way.