Inclusive fitness,asymmetric competition and kin selection in plants

The findings that some plants alter their competitive phenotype in response to genetic relatedness of its conspecific neighbour (and presumed competitor) has spurred an increasing interest in plant kin‐interactions. This phenotypic response suggests the ability to assess the genetic relatedness of conspecific competitors, proposing kin selection as a process that can influence plant competitive interactions. Kin selection can favour restrained competitive growth towards kin, if the fitness loss from reducing own growth is compensated by increased fitness in the related neighbour. This may lead to positive frequency dependency among related conspecifics with important ecological consequences for species assemblage and coexistence. However, kin selection in plants is still controversial. First, many studies documenting a plastic response to neighbour relatedness do not estimate fitness consequences of the individual that responds, and when estimated, fitness of individuals grown in competition with kin did not necessarily exceed that of individuals grown in non‐kin groups. Although higher fitness in kin groups could be consistent with kin selection, this could also arise from mechanisms like asymmetric competition in the non‐kin groups. Here we outline the main challenges for studying kin selection in plants taking genetic variation for competitive ability into account. We emphasize the need to measure inclusive fitness in order to assess whether kin selection occurs, and show under which circumstances kin selected responses can be expected. We also illustrate why direct fitness estimates of a focal plant, and group fitness estimates are not suitable for documenting kin selection. Importantly, natural selection occurs at the individual level and it is the inclusive fitness of an individual plant – not the mean fitness of the group – that can capture if a differential response to neighbour relatedness is favoured by kin selection.     

Evidence for competition and cooperation among climbing plants

A plant's best strategy for acquiring resources may often depend on the identity of neighbours. Here, I ask whether plants adjust their strategy to local relatedness: individuals may cooperate (reduce competitiveness) with kin but compete relatively intensely with non-kin. In a greenhouse experiment with Ipomoea hederacea, neighbouring siblings from the same inbred line were relatively uniform in height; groups of mixed lines, however, were increasingly variable as their mean height increased. The reproductive yield of mixed and sibling groups was similar overall, but when adjusted to a common mean height and height inequality, the yield of mixed groups was significantly less. Where this difference in yield was most pronounced (among groups that varied most in height), mixed groups tended to allocate more mass to roots than comparable sibling groups, and overall, mixed groups produced significantly fewer seeds per unit mass of roots. These results suggest that, from the group perspective, non-kin may have wasted resources in below-ground competition at the expense of reproduction; kin groups, on the other hand, displayed the relative efficiency that is expected of reduced competitiveness.     

Size inequality and the tragedy of the commons phenomenon in plant competition

Game theory predicts that the evolutionarily stable level of root production is greater for plants grown with neighbours compared to plants grown alone, even when the available resources per plant are constant. This follows from the fact that for plants grown alone, new roots compete only with other roots on the same plant, whereas for multiple plants grown in a group, new roots can also compete with the roots of other plants, thereby potentially acquiring otherwise unavailable resources at their neighbours’ expense. This phenomenon, which results in plants grown with neighbours over-proliferating roots at the expense of above-ground biomass, has been described as a ‘tragedy of the commons’, and requires that plants can distinguish self from non-self tissues. While this game theoretical model predicts the evolutionarily stable strategies of individual plants, it has only been tested on average allocation patterns of groups of plants. This is problematic, because average patterns can appear to reflect a tragedy of the commons, even when none has occurred. In particular, assuming (1) a decelerating relationship between individual plant biomass and the amount of resources available, and (2) greater size inequality in plants grown with neighbours compared to plants grown alone (due to asymmetric competition), then plants grown with neighbours should, at least on average, be smaller than plants grown alone. This is a manifestation of ‘Jensen’s Inequality’, which states that for decelerating functions, the average value of the function is less than the function of the average value. We suggest that Jensen’s Inequality should serve as an appropriate null hypothesis for examining biologically-based explanations of changes in biomass allocation strategies.     

The shift from plant–plant facilitation to competition under severe water deficit is spatially explicit

The stress‐gradient hypothesis predicts a higher frequency of facilitative interactions as resource limitation increases. Under severe resource limitation, it has been suggested that facilitation may revert to competition, and identifying the presence as well as determining the magnitude of this shift is important for predicting the effect of climate change on biodiversity and plant community dynamics. In this study, we perform a meta‐analysis to compare temporal differences of species diversity and productivity under a nurse plant (Retama sphaerocarpa) with varying annual rainfall quantity to test the effect of water limitation on facilitation. Furthermore, we assess spatial differences in the herbaceous community under nurse plants in situ during a year with below‐average rainfall. We found evidence that severe rainfall deficit reduced species diversity and plant productivity under nurse plants relative to open areas. Our results indicate that the switch from facilitation to competition in response to rainfall quantity is nonlinear. The magnitude of this switch depended on the aspect around the nurse plant. Hotter south aspects under nurse plants resulted in negative effects on beneficiary species, while the north aspect still showed facilitation. Combined, these results emphasize the importance of spatial heterogeneity under nurse plants for mediating species loss under reduced precipitation, as predicted by future climate change scenarios. However, the decreased water availability expected under climate change will likely reduce overall facilitation and limit the role of nurse plants as refugia, amplifying biodiversity loss.     

High densities of leaf‐tiers in open habitats are explained by host plant architecture

1. Crown architecture remains one of the least studied plant traits that influence plant–herbivore interactions. The hypotheses that dense crown architecture of mountain birches from open habitats favours leaf‐tying caterpillars through bottom‐up and/or top‐down effects associated with high leaf connectivity were tested. 2. Population densities of leaf‐tying herbivores in open (industrial barren and seashore) habitats were three times as high as in the shaded (forest) habitats. An experimental increase in leaf density by branch binding did not affect foliar consumption by free‐living herbivores but increased consumption by leaf‐tiers. 3. The specific leaf weight was lower in shaded habitats and in bound branches, but branch binding did not influence either the foliar concentrations of carbon and nitrogen or the pupal weight of the most abundant leaf‐tier, Carpatolechia proximella Hbn. (Lepidoptera: Gelechiidae). 4. Caterpillars of C. proximella build several shelters during their lifetime and spend a considerable amount of time outside the shelter, where they excrete most of their faeces. In bound branches, caterpillars built new shelters more frequently than in control branches, and consumed less foliar biomass per shelter. 5. Mortality from parasitoids in bound branches was half that in the control, presumably because the complex environment disrupted parasitoid searching behaviour and/or because of lower damage to leaves from which the shelters were built. 6. It is concluded that the crown architecture associated with high leaf connectivity decreases mortality risks from natural enemies both outside and inside the shelter. Compact and dense crowns of host plants may at least partly explain high population densities of leaf‐tiers in open habitats.