Peter Alpert

Invasiveness, invasibility and the role of environmental stress in the spread of non-native plants

Abstract Invasion ecology, the study of how organisms spread in habitats to which they are not native, asks both about the invasiveness of species and the invasibility of habitats: Which species are most likely to become invasive? Which habitats are most susceptible to invasion? To set the stage for considering these questions with regard to plants, we offer a two-way classification of nativeness and invasiveness that distinguishes natives, non-invasive non-natives and invasive non-natives. We then consider the current state of knowledge about invasiveness and invasibility. Despite much investigation, it has proven difficult to identify traits that consistently predict invasiveness. This may be largely because different traits favour invasiveness in different habitats. It has proven easier to identify types of habitats that are relatively invasible, such as islands and riverbanks. Factors thought to render habitats invasible include low intensities of competition, altered disturbance regimes and low levels of environmental stress, especially high resource availability. These factors probably often interact; the combination of altered disturbance with high resource availability may particularly promote invasibility. When biotic factors control invasibility, non-natives that are unlike native species may prove more invasive; the converse may also be true. We end with a simple conceptual model for cases in which high levels of environmental stress should and should not reduce invasibility. In some cases, it may be possible to manipulate stress to control biological invasions by plants.

The relative advantages of plasticity and fixity in different environments: when is it good for a plant to adjust?

Plant populations and species differ greatly in phenotypic plasticity. This could be because plasticity is advantageous under some conditions and disadvantageous or not advantageous under others. We distinguish adaptive from injurious and neutral plasticity and discuss when selection should favor adaptive plasticity over genetic differentiation or lack of phenotypic variation. It seems reasonable to hypothesize that selection is likely to favor plasticity when an environmental factor varies on the same spatial scale as the plant response unit, when the plant can respond to an environmental factor faster than the level of the factor changes, and when environmental variation is highly but not completely predictable. Phenotypic plasticity might also tend to be more advantageous when mean resource availability is high rather than low, when a response can occur late in development rather than early, and when a response is reversible rather than irreversible. There is substantial evidence for the hypothesis that predictability favors plasticity. However, available evidence does not support the hypothesis that high mean resource availability necessarily favors plasticity. Testing hypotheses about when it is good for a plant to adjust is central to understanding the diversity of plasticity in plants.

Desiccation-tolerance in bryophytes: a review

Abstract Desiccation-tolerance (DT), the ability to lose virtually all free intracellular water and then recover normal function upon rehydration, is one of the most remarkable features of bryophytes. The physiology of bryophytes differs in major respects from that of vascular plants by virtue of their smaller size; unlike vascular plants, the leafy shoots of bryophytes equilibrate rapidly with the water potential in their surroundings and tend to be either fully hydrated or desiccated and metabolically inactive. The time required to recover from desiccation increases and degree of recovery decreases with length of desiccation; both also depend upon temperature and intensity of desiccation. Tolerance in at least some species shows phenotypic plasticity. Recovery of respiration, photosynthesis and protein synthesis takes place within minutes or an hour or two; recovery of the cell cycle, food transport and the cytoskeleton may take 24 hours or more. Positive carbon balance is essential to survival of repeated cycles of drying and wetting; significant growth requires continuously wet periods of a few days or more. Male and female gametophytes, and gametophyte and sporophyte, may differ in tolerance. Desiccation-tolerance is essential to dispersal and establishment of spores and vegetative propagules. The mechanisms of DT in bryophytes, including expression of LEA proteins, high content of non-reducing sugars and effective antioxidant and photo-protection, are at least partly constitutive, allowing survival of rapid drying, but changes in gene expression resulting from mRNA sequestration and alterations in translational controls elicited upon rehydration are also important to repair processes following re-wetting. Phylogenetic and ecological considerations suggest that DT is a primitive character of land plants, lost in the course of evolution of the homoiohydric vascular-plant shoot system, but retained in spores, pollen and seeds, and re-evolved in the vegetative tissues of vascular “resurrection plants.” Bryophytes have retained the poikilohydry and DT that are probably the optimal pattern of adaptation at their scale, but modern bryophytes are specialized and diverse, and are removed by the same span of evolutionary time as the flowering plants from their primitive origins.

Constraints of tolerance: why are desiccation-tolerant organisms so small or rare?

SUMMARY Drying to equilibrium with the air kills nearly all animals and flowering plants, including livestock and crops. This makes drought a key ecological problem for terrestrial life and a major cause of human famine. However, the ability to tolerate complete desiccation is widespread in organisms that are either <5 mm long or found mainly where desiccation-sensitive organisms are scarce. This suggests that there is a trade-off between desiccation tolerance and growth. Recent molecular and biochemical research shows that organisms tolerate desiccation through a set of mechanisms, including sugars that replace water and form glasses, proteins that stabilize macromolecules and membranes, and anti-oxidants that counter damage by reactive oxygen species. These protections are often induced by drying, and some of the genes involved may be homologous in microbes, plants and animals. Understanding how mechanisms of desiccation tolerance may constrain growth might show how to undo the constraint in some economically important macroorganisms and elucidate the much-studied but elusive relationship between tolerance of stress and productivity.

The Limits and Frontiers of Desiccation-Tolerant Life

Abstract Drying to equilibrium with the air is lethal to most species of animals and plants, making drought (i.e., low external water potential) a central problem for terrestrial life and a major cause of agronomic failure and human famine. Surprisingly, a wide taxonomic variety of animals, microbes, and plants do tolerate complete desiccation, defined as water content below 0.1 g H2O g−1 dry mass. Species in five phyla of animals and four divisions of plants contain desiccation-tolerant adults, juveniles, seeds, or spores. There seem to be few inherent limits on desiccation tolerance, since tolerant organisms can survive extremely intense and prolonged desiccation. There seems to be little phylogenetic limitation of tolerance in plants but may be more in animals. Physical constraints may restrict tolerance of animals without rigid skeletons and to plants shorter than 3 m. Physiological constraints on tolerance in plants may include control by hormones with multiple effects that could link tolerance to slow growth. Tolerance tends to be lower in organisms from wetter habitats, and there may be selection against tolerance when water availability is high. Our current knowledge of limits to tolerance suggests that they pose few obstacles to engineering tolerance in prokaryotes and in isolated cells and tissues, and there has already been much success on this scientific frontier of desiccation tolerance. However, physical and physiological constraints and perhaps other limits may explain the lack of success in extending tolerance to whole, desiccation-sensitive, multicellular animals and plants. Deeper understanding of the limits to desiccation tolerance in living things may be needed to cross this next frontier.

Nitrogen sharing among ramets increases clonal growth in Fragaria chiloensis

Transport of resources among connected ramets can increase the growth of clonal plants when ramets are located in microsites with contrasting light, water, or total soil nutrient availability. To test whether this is also true for soil nitrogen availability alone, two greenhouse experiments were conducted using the stoloniferous herb Fragaria chi- loensis (beach strawberry). First, pairs of ramets were prepared in which the two ramets were connected by a stolon but rooted in separate pots. Ramets in a pair were: (1) severed from each other and given contrasting levels of soil nitrogen (high vs. low); (2) left connected and given contrasting levels of soil nitrogen; or (3) left connected and given a uniform level of soil nitrogen (both high or both low). Second, single ramets were given the high, the low, or one of three intermediate levels of soil nitrogen. Maintaining the vascular connection between ramets given contrasting levels of soil nitrogen resulted in a greater production of new stolons and new ramets by the ramet given the low nitrogen level, and in slightly greater leaf growth of the ramet given the low nitrogen level if it was also younger than the connected ramet. However, a ramet given high soil nitrogen grew much more than a ramet given low soil nitrogen, even when connected. Maintaining the vascular connection between ramets given a uniform level of soil nitrogen had no significant effect on growth. Single ramets given a higher soil nitrogen level had greater total and component masses, more stolons and new ramets, higher nitrogen concentrations, higher specific leaf area, and a higher proportional mass of leaves and of stolons and new ramets. Experiments suggested that: (1) transport of nitrogen among ramets can increase growth in F. chiloensis, but to a limited extent; (2) transport of nitrogen through a stolon may be either acropetal or basipetal but tends to promote the growth of younger ramets; (3) higher soil nitrogen availability alters plant architecture by increasing allocation to leaves and to new stolons and ramets; and (4) physiological integration among ramets modifies clonal architecture because nitro- gen acquired by roots promotes growth of both a rooted ramet and its new ramets, whereas nitrogen acquired from another ramet promotes the growth of new ramets almost exclu- sively.

Clonal integration in Fragaria chiloensis differs between populations: ramets from grassland are selfish

Abstract In plants, only species with clonal growth are able to directly transfer resources between otherwise independent units of the same genetic individual. A simple conceptual model of plant performance as a function of internal resource supply and environmental resource availability suggests that resource sharing between ramets within clones is likely to be disadvantageous in uniform habitats and advantageous in patchy ones. It was therefore hypothesized that clones in populations from relatively uniform habitats will have been selected for low rates of resource sharing between ramets compared to clones in populations from patchier habitats. In coastal northern California, the clonal herb Fragaria chiloensis is common both in grasslands, where resources are relatively uniform, and on sand dunes, where resources are more patchy. It was predicted that clones from a grassland population of Fragaria would have “selfish” ramets with low rates of resource sharing compared to clones from an adjacent dune population. Ramets were subjected to contrasting light levels with and without connection between ramets. Patterns of biomass accumulation were consistent with the prediction. This appears to be the first report of genetically based variation in patterns of resource sharing in clonal plants. It supports the idea that these patterns are locally selected to increase plant performance in habitats with different patterns of resource availability.

Integrated Conservation and Development Projects Examples from Africa

ntegrated conservation and development projects, or ICDPs, represent a new approach to the conservation of biodiversity and ecological systems in developing countries (Wells and Brandon 1993). ICDPs distinguish themselves from other approaches by setting a dual and equal focus on biological conservation and human development. Their main goal is to link conservation and development such that each fosters the other. Even though conservation and development have generally figured as antithetical alternatives, ICDPs have multiplied throughout the developing world in little more than a decade. More than 100 ICDPs have been described (Alpert 1993, Anderson and Grove 1987, BSP 1993, Butynski and Kalina 1993, Goldstein 1994, Hamilton et al. 1993, Lewis and Carter 1993, Lucas 1992, McNeely 1993, 1995, Potter et al. 1993, Redford and Padoch 1992, Wells et al. 1992, West and Brechin 1991, Western et al. 1994, Wright 1992), including more than 50 in at least 20 countries of sub-Saharan Africa. Earlier reviews have judged ICDPs to be promising but unproven (BranICDPs aim to fill the

RECIPROCAL TRANSPORT BETWEEN RAMETS INCREASES GROWTH OF FRAGARIA CHILOENSIS WHEN LIGHT AND NITROGEN OCCUR IN SEPARATE PATCHES BUT ONLY IF PATCHES ARE RICH

SummaryFragaria chiloensis is a stoloniferous perennial herb that grows on coastal sand dunes where scattered shrubs create small patches of lower photon flux density (PFD) but higher soil nitrogen availability. The potential effects of resource transport between ramets when PFD and soil nitrogen are negatively associated in space were tested by comparing the growth of pairs of ramets in which the vascular connection between ramets was either severed or left intact. One ramet in each pair was given high PFD but a low level of soil nitrogen and the other ramet was given low PFD but high N. The analogous effects of resource transport likely to be realized in nature were tested by substituting a more realistic medium soil nitrogen level in place of the high level. Results suggested that connected ramets exchanged carbon and nitrogen under both regimes of soil nitrogen heterogeneity. In the low versus high nitrogen regime, connected ramets had higher combined dry biomass and different patterns of dry mass partitioning from those of severed ramets; effect of connection was greater on ramets given low PFD and high N and on younger ramets. In the low versus medium nitrogen regime, connected ramets had different patterns of partitioning only. Apparent reciprocal resource transport between ramets can enhance the growth of ramets with complementary resource deficiencies, but may affect growth in dry mass only when maximum resource levels are high.

Root cooperation in a clonal plant: connected strawberries segregate roots.

Abstract. The ability to selectively avoid competition with members of the same clone should be highly advantageous but has not been demonstrated in plants. We found that physical connection between plants in a clone of the wild strawberry Fragaria chiloensis induced them to segregate their roots, significantly increasing clonal performance. Such increase in performance was not found when plants were grown in containers that artificially divided their rooting zones. There was no effect of connection in a different clone of F. chiloensis with a lower degree of carbon transport between connected plants, suggesting that the mechanism for root segregation depended upon transport of a signal through the strawberry runners. We suggest that clonal integration allows some clones to coordinate below-ground resource foraging with other clone members, thus exhibiting a type of root cooperation.