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Mikko Heino

Publications and source records attributed to Mikko Heino.

9 recordsLinked to original sources

Climate change and condition of herring (Clupea harengus) explain long-term trends in extent of skipped reproduction.

It is commonly assumed that iteroparous fish, once mature, normally reproduce in all consecutive seasons. Recent work has suggested, however, that in Norwegian spring-spawning herring-a population that undertakes extensive spawning migrations-almost one in two adults may skip their second spawning migration. Why should herring not return to spawn the year after first spawning, but instead wait an extra year? For herring, participation in distant, energetically costly, and risky spawning migrations will only pay off in terms of fitness if individuals are sufficiently large, and in sufficient condition, to both successfully migrate and spawn. Changes in the environment and individual condition should therefore affect the likelihood of skipped spawning. This paper describes long-term changes in the extent to which the second reproductive season is skipped in this herring population. These are shown to be linked to the size and condition of herring as first-time spawners, and to climatic factors possibly related to food availability. The findings corroborate the hypothesis that skipped reproduction results from trade-offs between current and future reproduction, growth and survival.

Animals↗

The adaptive dynamics of function-valued traits.

This study extends the framework of adaptive dynamics to function-valued traits. Such adaptive traits naturally arise in a great variety of settings: variable or heterogeneous environments, age-structured populations, phenotypic plasticity, patterns of growth and form, resource gradients, and in many other areas of evolutionary ecology. Adaptive dynamics theory allows analysing the long-term evolution of such traits under the density-dependent and frequency-dependent selection pressures resulting from feedback between evolving populations and their ecological environment. Starting from individual-based considerations, we derive equations describing the expected dynamics of a function-valued trait in asexually reproducing populations under mutation-limited evolution, thus generalizing the canonical equation of adaptive dynamics to function-valued traits. We explain in detail how to account for various kinds of evolutionary constraints on the adaptive dynamics of function-valued traits. To illustrate the utility of our approach, we present applications to two specific examples that address, respectively, the evolution of metabolic investment strategies along resource gradients, and the evolution of seasonal flowering schedules in temporally varying environments.

Adaptation, Biological↗

Function-valued adaptive dynamics and the calculus of variations.

Adaptive dynamics has been widely used to study the evolution of scalar-valued, and occasionally vector-valued, strategies in ecologically realistic models. In many ecological situations, however, evolving strategies are best described as function-valued, and thus infinite-dimensional, traits. So far, such evolution has only been studied sporadically, mostly based on quantitative genetics models with limited ecological realism. In this article we show how to apply the calculus of variations to find evolutionarily singular strategies of function-valued adaptive dynamics: such a strategy has to satisfy Euler's equation with environmental feedback. We also demonstrate how second-order derivatives can be used to investigate whether or not a function-valued singular strategy is evolutionarily stable. We illustrate our approach by presenting several worked examples.

Biological Evolution↗

Scale analysis suggests frequent skipping of the second reproductive season in Atlantic herring.

A common assumption in fish studies is that iteroparous fishes, once mature, normally reproduce in all consecutive seasons. An analysis of scales from Norwegian spring-spawning herring collected between 1935 and 1973 revealed a strong under-representation (47% of expected) of second-time spawners on the spawning grounds. This reduction is not satisfactorily explained by errors in scale-reading, suggesting that the second reproductive season is frequently skipped. Skipping a season may relate to trade-offs between growth, current and future reproduction, and survival, which are likely to be particularly strong for young adult herring.

Age Factors↗

Maturation trends indicative of rapid evolution preceded the collapse of northern cod.

Northern cod, comprising populations of Atlantic cod (Gadus morhua) off southern Labrador and eastern Newfoundland, supported major fisheries for hundreds of years. But in the late 1980s and early 1990s, northern cod underwent one of the worst collapses in the history of fisheries. The Canadian government closed the directed fishing for northern cod in July 1992, but even after a decade-long offshore moratorium, population sizes remain historically low. Here we show that, up until the moratorium, the life history of northern cod continually shifted towards maturation at earlier ages and smaller sizes. Because confounding effects of mortality changes and growth-mediated phenotypic plasticity are accounted for in our analyses, this finding strongly suggests fisheries-induced evolution of maturation patterns in the direction predicted by theory. We propose that fisheries managers could use the method described here as a tool to provide warning signals about changes in life history before more overt evidence of population decline becomes manifest.

Aging↗

Adaptive changes in harvested populations: plasticity and evolution of age and size at maturation.

We investigate harvest-induced adaptive changes in age and size at maturation by modelling both plastic variation and evolutionary trajectories. Harvesting mature individuals displaces the reaction norm for age and size at maturation toward older ages and larger sizes and rotates it clockwise, whereas harvesting immature individuals has the reverse qualitative effect. If both immature and mature individuals are harvested, the net effect has approximately the same trend as when harvesting immature individuals only. This stems from the sensitivity of the evolutionary response, which depends on the maturity state of harvested individuals, but also on the type of harvest mortality (negatively or positively density dependent, density independent) and the value of three life-history parameters (natural mortality, growth rate and the trade-off between growth and reproduction). Evolutionary changes in the maturation reaction norm have strong repercussions for the mean size and the density of harvested individuals that, in most cases, result in the reduction of biomass--a response that population dynamical models would overlook. These results highlight the importance of accounting for evolutionary trends in the long-term management of exploited living resources and give qualitative insights into how to minimize the detrimental consequences of harvest-induced evolutionary changes in maturation reaction norms.

Adaptation, Biological↗

Unexpected discontinuities in life-history evolution under size-dependent mortality.

In many organisms survival depends on body size. We investigate the implications of size-selective mortality on life-history evolution by introducing and analysing a new and particularly flexible life-history model with the following key features: the lengths of growth and reproductive periods in successive reproductive cycles can vary evolutionarily, the model does not constrain evolution to patterns of either determinate or indeterminate growth, and lifetime number and sizes of broods are the outcomes of evolutionarily optimal life-history decisions. We find that small changes in environmental conditions can lead to abrupt transitions in optimal life histories when size-dependent mortality is sufficiently strong. Such discontinuous switching results from antagonistic selection pressures and occurs between strategies of early maturation with short reproductive periods and late maturation with long reproductive cycles. When mortality is size-selective and the size-independent component is not too high, selection favours prolonged juvenile growth, thereby allowing individuals to reach a mortality refuge at large body size before the onset of reproduction. When either component of mortality is then increased, the mortality refuge first becomes unattractive and eventually closes up altogether, resulting in short juvenile growth and frequent reproduction. Our results suggest a new mechanism for the evolution of life-history dimorphisms.

Algorithms↗

Metapopulation-level adaptation of insect host plant preference and extinction-colonization dynamics in heterogeneous landscapes.

Species living in highly fragmented landscapes typically occur as metapopulations with frequent turnover of local populations. The turnover rate depends on population sizes and connectivities, but it may also depend on the phenotypic and genotypic composition of populations. The Glanville fritillary butterfly (Melitaea cinxia) in Finland uses two host plant species, which show variation in their relative abundances at two spatial scales: locally among individual habitat patches and regionally among networks of patches. Female butterflies in turn exhibit spatial variation in genetically determined host plant preference within and among patch networks. Emigration, immigration and establishment of new populations have all been shown to be strongly influenced by the match between the host plant composition of otherwise suitable habitat patches and the host plant preference of migrating butterflies. The evolutionary consequences of such biased migration and colonization with respect to butterfly phenotypes might differ depending on spatial configuration and plant species composition of the patches in heterogeneous patch networks. Using a spatially realistic individual-based model we show that the model-predicted evolution of host plant preference due to biased migration explains a significant amount of the observed variation in host plant use among metapopulations living in dissimilar networks. This example illustrates how the ecological extinction-colonization dynamics may be linked with the evolutionary dynamics of life history traits in metapopulations.

Adaptation, Physiological↗

Measuring probabilistic reaction norms for age and size at maturation.

We present a new probabilistic concept of reaction norms for age and size at maturation that is applicable when observations are carried out at discrete time intervals. This approach can also be used to estimate reaction norms for age and size at metamorphosis or at other ontogenetic transitions. Such estimations are critical for understanding phenotypic plasticity and life-history changes in variable environments, assessing genetic changes in the presence of phenotypic plasticity, and calibrating size- and age-structured population models. We show that previous approaches to this problem, based on regressing size against age at maturation, give results that are systematically biased when compared to the probabilistic reaction norms. The bias can be substantial and is likely to lead to qualitatively incorrect conclusions; it is caused by failing to account for the probabilistic nature of the maturation process. We explain why, instead, robust estimations of maturation reaction norms should be based on logistic regression or on other statistical models that treat the probability of maturing as a dependent variable. We demonstrate the utility of our approach with two examples. First, the analysis of data generated for a known reaction norm highlights some crucial limitations of previous approaches. Second, application to the northeast arctic cod (Gadus morhua) illustrates how our approach can be used to shed new light on existing real-world data.

Age Factors↗