Izza Ghani | University of Helsinki (original) (raw)

Papers by Izza Ghani

Research paper thumbnail of Evidence for genetic differentiation in timing of maturation among nine‐spined stickleback populations

Journal of Evolutionary …, 2013

Timing of maturation is an important life-history trait that is likely to be subjected to strong ... more Timing of maturation is an important life-history trait that is likely to be subjected to strong natural selection. Although population differences in timing of maturation have been frequently reported in studies of wild animal populations, little is known about the genetic basis of this differentiation. Here, we investigated population and sex differences in timing of maturation within and between two nine-spined stickleback (Pungitius pungitius) populations in a laboratory breeding experiment. We found that fish from the high-predation marine population matured earlier than fish from the low-predation pond population and males matured earlier than females. Timing of maturation in both reciprocal hybrid crosses between the two populations was similar to that in the marine population, suggesting that early timing of maturation is a dominant trait, whereas delayed timing of maturation in the pond is a recessive trait. Thus, the observed population divergence is suggestive of strong natural selection against early maturation in the piscine-predator-free pond population.

Research paper thumbnail of Evolution of stickleback feeding behaviour: genetics of population divergence at different ontogenetic stages

Journal of Evolutionary …, 2013

The evolutionary significance of individual consistency in a given behaviour called animal person... more The evolutionary significance of individual consistency in a given behaviour called animal personalityhas been subject to a lot of recent research. However, the genetic underpinnings of population divergence in mean personality have rarely been studied, especially across different ontogenetic stages. Previous work has shown that marine vs. pond populations of ninespined sticklebacks (Pungitius pungitius) have undergone adaptive divergence in a series of fitness-related traits, including behaviour. One particular behavioural trait important in this system is feeding activity: giant pond sticklebacks are more active feeders than their normal sized marine conspecifics. In a common garden experiment, we raised individuals from pure and hybrid F 1 -generation crosses of a highly divergent marinepond population pair to see if (i) feeding activity and/or its ontogenetic change was consistent between individuals, and if (ii) population divergence at different ontogenetic stages could be explained by additive genetic, nonadditive genetic or maternal effects. We found that feeding activity decreased with age, but that these changes were consistently different among both individuals and crosses. The among cross patterns were consistent with a nonadditive genetic scenario: in the early period pond sticklebacks expressed dominance for high feeding activity, while in the late period marine sticklebacks expressed dominance for low feeding activity. We conclude that nine-spined sticklebacks exhibit different feeding personalities, and that the population divergence in feeding personality is explainable by age-dependent expression of genetic dominance.

Research paper thumbnail of Contrasting population structures in two sympatric fishes in the Baltic Sea basin

Marine biology, 2012

Detailed multispecies studies on the patterns of genetic variability and differentiation in marin... more Detailed multispecies studies on the patterns of genetic variability and differentiation in marine environments are still rare. Using mitochondrial and nuclear genetic markers, we compared genetic variability and population structuring of threespine (Gasterosteus aculeatus) and ninespine (Pungitius pungitius) sticklebacks from the same eleven marine and six freshwater locations within the Baltic Sea basin. Analyses of both marker types revealed a significantly lower degree of genetic structuring in both marine and freshwater populations of threespine than those ninespine sticklebacks. Isolation-by-distance (IBD) was detected across the marine populations in both species, suggesting spatially limited gene flow. However, the levels of genetic diversity and differentiation across the localities were uncorrelated between the two species in both marine and freshwater environments. Accordingly, estimates of effective population sizes were larger and migration rates were higher for three-than for ninespine sticklebacks. Hence, ninespine stickleback populations from the Baltic Sea basin appear to be subject to stronger genetic drift than sympatric threespine sticklebacks, and the proximate reason for this difference is likely to be found from autecological differences between the two species. In accordance with the earlier studies, genetic variability was higher and the degree of genetic differentiation was lower in marine than in freshwater populations in both species.

Research paper thumbnail of Body size divergence in nine‐spined sticklebacks: disentangling additive genetic and maternal effects

Biological Journal of the …, 2012

Interpopulation differences in body size are of common occurrence in vertebrates, but the relativ... more Interpopulation differences in body size are of common occurrence in vertebrates, but the relative importance of genetic, maternal, and environmental effects as causes of observed differentiation have seldom been assessed in the wild. Gigantism in pond nine-spined sticklebacks (Pungitius pungitius Linnaeus, 1758) has been repeatedly observed, but the quantitative genetic basis of population divergence in size has remained unstudied. We conducted a common garden experiment -using 'pure' and reciprocal crosses between two populations ('giant' pond versus 'normal' marine) -to test for the relative importance of additive genetic, non-additive genetic, and maternal effects on body size after 11 months of growth in the laboratory. We found that body size difference between the two populations in laboratory conditions owed mainly to additive genetic effects, and only to a minor degree to maternal effects. Furthermore, the weak maternal effects were seen only in the offspring of 'giant' mothers, and appeared to be mediated through differences in egg size. Thus, the results suggest that gigantism in pond populations of P. pungitius is based on the effects of additively acting genes, rather than to direct environmental induction, or maternal or non-additive gene action.

Research paper thumbnail of High levels of fluctuating asymmetry in isolated stickleback populations

BMC Evolutionary …, 2012

Background: Fluctuating asymmetry (FA), defined as small random deviations from the ideal bilater... more Background: Fluctuating asymmetry (FA), defined as small random deviations from the ideal bilateral symmetry, has been hypothesized to increase in response to both genetic and environmental stress experienced by a population. We compared levels of FA in 12 bilateral meristic traits (viz. lateral-line system neuromasts and lateral plates), and heterozygosity in 23 microsatellite loci, among four marine (high piscine predation risk) and four pond (zero piscine predation risk) populations of nine-spined sticklebacks (Pungitius pungitius). Results: Pond sticklebacks had on average three times higher levels of FA than marine fish and this difference was highly significant. Heterozygosity in microsatellite markers was on average two times lower in pond (H E % 0.3) than in marine (H E % 0.6) populations, and levels of FA and heterozygosity were negatively correlated across populations. However, after controlling for habitat effect on heterozygosity, levels of FA and heterozygosity were uncorrelated. Conclusions: The fact that levels of FA in traits likely to be important in the context of predator evasion were elevated in ponds compared to marine populations suggests that relaxed selection for homeostasis in ponds lacking predatory fish may be responsible for the observed habitat difference in levels of FA. This inference also aligns with the observation that the levels of genetic variability across the populations did not explain population differences in levels of FA after correcting for habitat effect. Hence, while differences in strength of selection, rather than in the degree of genetic stress could be argued to explain habitat differences in levels of FA, the hypothesis that increased FA in ponds is caused by genetic stress cannot be rejected.

Research paper thumbnail of Intraspecific divergence in the lateral line system in the nine‐spined stickleback (Pungitius pungitius)

Journal of …, 2011

The mechanosensory lateral line system of fishes is an important organ system conveying informati... more The mechanosensory lateral line system of fishes is an important organ system conveying information crucial to individual fitness. Yet, our knowledge of lateral line diversity is almost exclusively based on interspecific studies, whereas intraspecific variability and possible population divergence have remained largely unexplored. We investigated lateral line system variability in four marine and five pond populations of nine-spined stickleback (Pungitius pungitius). We found significant differences in neuromast number between pond and marine fish. In particular, three of seventeen lateral line regions (viz. caudal peduncle superficial neuromasts; canal neuromasts from the anterior trunk and caudal peduncle) showed strong divergence between habitats. Similar results were obtained with laboratory-reared individuals from a subset of populations, suggesting that the patterns found in nature likely have a genetic basis. Interestingly, we also found habitat-dependent population divergence in neuromast variability, with pond populations showing greater heterogeneity than marine populations, although only in wild-caught fish. A comparison of neutral genetic (F ST ) and phenotypic (P ST ) differentiation suggested that natural selection is likely associated with habitat-dependent divergence in neuromast counts. Hence, the results align with the conclusion that the mechanosensory lateral line system divergence among marine and pond nine-spined sticklebacks is adaptive.

Research paper thumbnail of Evidence for genetic differentiation in timing of maturation among nine‐spined stickleback populations

Journal of Evolutionary …, 2013

Timing of maturation is an important life-history trait that is likely to be subjected to strong ... more Timing of maturation is an important life-history trait that is likely to be subjected to strong natural selection. Although population differences in timing of maturation have been frequently reported in studies of wild animal populations, little is known about the genetic basis of this differentiation. Here, we investigated population and sex differences in timing of maturation within and between two nine-spined stickleback (Pungitius pungitius) populations in a laboratory breeding experiment. We found that fish from the high-predation marine population matured earlier than fish from the low-predation pond population and males matured earlier than females. Timing of maturation in both reciprocal hybrid crosses between the two populations was similar to that in the marine population, suggesting that early timing of maturation is a dominant trait, whereas delayed timing of maturation in the pond is a recessive trait. Thus, the observed population divergence is suggestive of strong natural selection against early maturation in the piscine-predator-free pond population.

Research paper thumbnail of Evolution of stickleback feeding behaviour: genetics of population divergence at different ontogenetic stages

Journal of Evolutionary …, 2013

The evolutionary significance of individual consistency in a given behaviour called animal person... more The evolutionary significance of individual consistency in a given behaviour called animal personalityhas been subject to a lot of recent research. However, the genetic underpinnings of population divergence in mean personality have rarely been studied, especially across different ontogenetic stages. Previous work has shown that marine vs. pond populations of ninespined sticklebacks (Pungitius pungitius) have undergone adaptive divergence in a series of fitness-related traits, including behaviour. One particular behavioural trait important in this system is feeding activity: giant pond sticklebacks are more active feeders than their normal sized marine conspecifics. In a common garden experiment, we raised individuals from pure and hybrid F 1 -generation crosses of a highly divergent marinepond population pair to see if (i) feeding activity and/or its ontogenetic change was consistent between individuals, and if (ii) population divergence at different ontogenetic stages could be explained by additive genetic, nonadditive genetic or maternal effects. We found that feeding activity decreased with age, but that these changes were consistently different among both individuals and crosses. The among cross patterns were consistent with a nonadditive genetic scenario: in the early period pond sticklebacks expressed dominance for high feeding activity, while in the late period marine sticklebacks expressed dominance for low feeding activity. We conclude that nine-spined sticklebacks exhibit different feeding personalities, and that the population divergence in feeding personality is explainable by age-dependent expression of genetic dominance.

Research paper thumbnail of Contrasting population structures in two sympatric fishes in the Baltic Sea basin

Marine biology, 2012

Detailed multispecies studies on the patterns of genetic variability and differentiation in marin... more Detailed multispecies studies on the patterns of genetic variability and differentiation in marine environments are still rare. Using mitochondrial and nuclear genetic markers, we compared genetic variability and population structuring of threespine (Gasterosteus aculeatus) and ninespine (Pungitius pungitius) sticklebacks from the same eleven marine and six freshwater locations within the Baltic Sea basin. Analyses of both marker types revealed a significantly lower degree of genetic structuring in both marine and freshwater populations of threespine than those ninespine sticklebacks. Isolation-by-distance (IBD) was detected across the marine populations in both species, suggesting spatially limited gene flow. However, the levels of genetic diversity and differentiation across the localities were uncorrelated between the two species in both marine and freshwater environments. Accordingly, estimates of effective population sizes were larger and migration rates were higher for three-than for ninespine sticklebacks. Hence, ninespine stickleback populations from the Baltic Sea basin appear to be subject to stronger genetic drift than sympatric threespine sticklebacks, and the proximate reason for this difference is likely to be found from autecological differences between the two species. In accordance with the earlier studies, genetic variability was higher and the degree of genetic differentiation was lower in marine than in freshwater populations in both species.

Research paper thumbnail of Body size divergence in nine‐spined sticklebacks: disentangling additive genetic and maternal effects

Biological Journal of the …, 2012

Interpopulation differences in body size are of common occurrence in vertebrates, but the relativ... more Interpopulation differences in body size are of common occurrence in vertebrates, but the relative importance of genetic, maternal, and environmental effects as causes of observed differentiation have seldom been assessed in the wild. Gigantism in pond nine-spined sticklebacks (Pungitius pungitius Linnaeus, 1758) has been repeatedly observed, but the quantitative genetic basis of population divergence in size has remained unstudied. We conducted a common garden experiment -using 'pure' and reciprocal crosses between two populations ('giant' pond versus 'normal' marine) -to test for the relative importance of additive genetic, non-additive genetic, and maternal effects on body size after 11 months of growth in the laboratory. We found that body size difference between the two populations in laboratory conditions owed mainly to additive genetic effects, and only to a minor degree to maternal effects. Furthermore, the weak maternal effects were seen only in the offspring of 'giant' mothers, and appeared to be mediated through differences in egg size. Thus, the results suggest that gigantism in pond populations of P. pungitius is based on the effects of additively acting genes, rather than to direct environmental induction, or maternal or non-additive gene action.

Research paper thumbnail of High levels of fluctuating asymmetry in isolated stickleback populations

BMC Evolutionary …, 2012

Background: Fluctuating asymmetry (FA), defined as small random deviations from the ideal bilater... more Background: Fluctuating asymmetry (FA), defined as small random deviations from the ideal bilateral symmetry, has been hypothesized to increase in response to both genetic and environmental stress experienced by a population. We compared levels of FA in 12 bilateral meristic traits (viz. lateral-line system neuromasts and lateral plates), and heterozygosity in 23 microsatellite loci, among four marine (high piscine predation risk) and four pond (zero piscine predation risk) populations of nine-spined sticklebacks (Pungitius pungitius). Results: Pond sticklebacks had on average three times higher levels of FA than marine fish and this difference was highly significant. Heterozygosity in microsatellite markers was on average two times lower in pond (H E % 0.3) than in marine (H E % 0.6) populations, and levels of FA and heterozygosity were negatively correlated across populations. However, after controlling for habitat effect on heterozygosity, levels of FA and heterozygosity were uncorrelated. Conclusions: The fact that levels of FA in traits likely to be important in the context of predator evasion were elevated in ponds compared to marine populations suggests that relaxed selection for homeostasis in ponds lacking predatory fish may be responsible for the observed habitat difference in levels of FA. This inference also aligns with the observation that the levels of genetic variability across the populations did not explain population differences in levels of FA after correcting for habitat effect. Hence, while differences in strength of selection, rather than in the degree of genetic stress could be argued to explain habitat differences in levels of FA, the hypothesis that increased FA in ponds is caused by genetic stress cannot be rejected.

Research paper thumbnail of Intraspecific divergence in the lateral line system in the nine‐spined stickleback (Pungitius pungitius)

Journal of …, 2011

The mechanosensory lateral line system of fishes is an important organ system conveying informati... more The mechanosensory lateral line system of fishes is an important organ system conveying information crucial to individual fitness. Yet, our knowledge of lateral line diversity is almost exclusively based on interspecific studies, whereas intraspecific variability and possible population divergence have remained largely unexplored. We investigated lateral line system variability in four marine and five pond populations of nine-spined stickleback (Pungitius pungitius). We found significant differences in neuromast number between pond and marine fish. In particular, three of seventeen lateral line regions (viz. caudal peduncle superficial neuromasts; canal neuromasts from the anterior trunk and caudal peduncle) showed strong divergence between habitats. Similar results were obtained with laboratory-reared individuals from a subset of populations, suggesting that the patterns found in nature likely have a genetic basis. Interestingly, we also found habitat-dependent population divergence in neuromast variability, with pond populations showing greater heterogeneity than marine populations, although only in wild-caught fish. A comparison of neutral genetic (F ST ) and phenotypic (P ST ) differentiation suggested that natural selection is likely associated with habitat-dependent divergence in neuromast counts. Hence, the results align with the conclusion that the mechanosensory lateral line system divergence among marine and pond nine-spined sticklebacks is adaptive.