Covariation among floral traits in Spergularia marina (Caryophyllaceae): geographic and temporal variation in phenotypic and among-family correlations

Strong covariation among traits suggests the presence of constraints on their independent evolution due to pleiotropy, to linkage, or to selective forces that maintain particular trait combinations. We examined floral trait covariation among individuals, among maternal families within and across populations, and over time, in greenhouse-raised plants of the autogamous Spergularia marina. We had three aims. First, since the phenotype of traits expressed by modular organs often changes as individuals age, estimates of the degree of genetic covariation between such traits may also change over time. To seek evidence for this, we measured weekly (for five weeks) an array of floral traits among plants representing ～ 10 maternal families from each of four populations. The statistical significance of the phenotypic and among-family correlations among traits changed over time. Second, we compared populations with respect to trait covariation to determine whether populations or traits appear to be evolving independently of one another. Differences observed among populations suggest that they have diverged genetically. Third, we sought correlations that might reflect constraints on the independent evolution of floral traits. Investment in another and ovule production per flower vary independently among maternal families; there was no evidence for a “trade-off” between male and female investment. We propose that in autogamous taxa one should not find a negative correlation between pollen and ovule production per flower, as such taxa cannot evolve sexual specialization and should be under strong selection to maintain an efficient pollen:ovule ratio, preventing the evolution of male-biased or female-biased genotypes. We found that other pairs of floral traits, however, expressed highly signficant correlation coefficients, suggesting the presence of some evolutionary constraints, at least within some populations, although their strength depended on exactly when flowers were sampled.     

Trophic links,nutrient fluxes,and natural history in the Allium ursinum food web,with particular reference to life history traits of two hoverfly herbivores (Diptera: Syrphidae)

The food web centering on Allium ursinum (Liliaceae) in a beech forest (Germany) is described, and temporal variation of active trophic links is related to species' life cycles. The most important insect herbivores are Cheilosia fasciata (a larval leaf miner) and Portevinia maculata (a larva bulb miner) (Diptera: Syrphidae). Energy, carbon and nitrogen flow in the food chain (Allium-Cheilosia-Phygadeuon ursini) are investigated and analysed with respect to differences in resource allocation by the leaf miner and its hymenoptereous parasitoid. In C. fasciata nitrogen is likely to be the limiting resource, while growth in Phygadeuon ursini appears energy-limited. Larval feeding habits of C. fasciata and Portevinia maculata determined the timing of the species' life cycles and, as a consequence, appeared to preclude the existence of a pupal parasitoid in Portevinia maculata. Further details of life history traits are demonstrated and discussed.     

Root to shoot ratio of crops as influenced by CO2

Crops of tomorrow are likely to grow under higher levels of atmospheric CO₂. Fundamental crop growth processes will be affected and chief among these is carbon allocation. The root to shoot ratio (R:S, defined as dry weight of root biomass divided by dry weight of shoot biomass) depends upon the partitioning of photosynthate which may be influenced by environmental stimuli. Exposure of plant canopies to high CO₂ concentration often stimulates the growth of both shoot and root, but the question remains whether elevated atmospheric CO₂ concentration will affect roots and shoots of crop plants proportionally. Since elevated CO₂ can induce changes in plant structure and function, there may be differences in allocation between root and shoot, at least under some conditions. The effect of elevated atmospheric CO₂ on carbon allocation has yet to be fully elucidated, especially in the context of changing resource availability. Herein we review root to shoot allocation as affected by increased concentrations of atmospheric CO₂ and provide recommendations for further research. Review of the available literature shows substantial variation in R:S response for crop plants. In many cases (59.5%) R:S increased, in a very few (3.0%) remained unchanged, and in others (37.5%) decreased. The explanation for these differences probably resides in crop type, resource supply, and other experimental factors. Efforts to understand allocation under CO₂ enrichment will add substantially to the global change response data base.Abbreviations R:S root to shoot ratio, dry weight basis     

Seasonal variation in growth, mortality and fat stores of roach and perch in Lough Neagh, Northern Ireland

The annual cycle for roach ( Rutilus rutilus ) and perch ( Perca fluviatilis ) in Lough Neagh, Northern Ireland can be divided into warm 'growing' and cold 'non-growing' seasons. Summer water temperatures declined between 1991–1993, as did the sizes of 0+ fish of both species. Small overwinter increases in length occurred after poor summers due, in roach, to size-selective mortality. Fat content was size-dependent and increased over the summer in both species. All sizes of roach but only small perch lost fat overwinter. Dead roach collected towards the end of the 1992/3 winter had significantly less fat than live individuals. The allocation of energy to growth (and ultimately reproduction) and to fat (and overwinter survival) was modelled as a tradeoff, assuming that fat has twice the energy density of protein and using observed summer and winter changes in body composition. The predicted allocation optima were close to observed values but, for roach, this entailed a substantial survivorship cost.     

OVULE PACKAGING IN STOCHASTIC POLLINATION AND FERTILIZATION ENVIRONMENTS

The modular morphology of plants has important consequences for reproductive strategies. Ovules are packaged in discrete structures (flowers) that usually vary stochastically in pollen capture and ovule fertilization, because of the vagaries of pollen transfer by external agents. Different ovule packaging schemes may use limited reproductive resources more or less effectively, so that some number of ovules per flower may be optimal, given the prevailing probabilities of ovule fertilization. I derive a phenotypic model for ovule number per flower that maximizes the expected total ovule fertilizations on a plant when pollination and fertilization vary randomly among individual flowers. This model predicts that, except for small or inexpensive flowers, ovules should be “oversupplied” relative to the mean receipt of pollen tubes, so that pollen limitation of seed set should be common. Published data are congruent with this prediction. Additional hypotheses on the relation of ovule packaging to floral cost, plant size, and variance in pollen receipt are suggested by the model, but few data exist to evaluate these hypotheses.     

Effects of litter accumulation and water table on morphology and productivity of Phragmites australis

Phragmites australis (Common reed) occurs in the interface between water and land. The water depth gradient from deep water to dry land is inversely related to litter accumulation. Eutrophication can result in an excessive production of litter, which may have a large impact on the occurrence of P. australis in this gradient. In an outdoor pot experiment, it was investigated how water tables in combination with substrates containing variable amounts of litter affect morphology and productivity of P. australis. Vegetatively propagated P. australis was grown in pots filled with river sand, litter, and different mixtures of sand and litter (25, 50 and 75% by volume). Four water table treatments were applied; drained (–12 cm), waterlogged (0 cm), flooded (+12 cm), and weekly fluctuating drained and flooded conditions (–12/+12 cm of water relative to substrate level). When drained, no differences between substrate treatments were present. Waterlogging, flooding fluctuating water table treatments caused growth reduction in substrate containing litter. The plants formed short shoots and thin rhizomes. With increasing water table, allocation of dry matter to stems increased at the expense of leaves and rhizomes. At intermediate levels of litter in the substrate, allocation to leaves was lowest. In both instances a lower leaf weight ratio (LWR) was (partly) compensated for by a higher specific leaf area (SLA), resulting in less pronounced differences in leaf area ratio (LAR). Aquatic roots developed when plants were waterlogged or flooded, and increased when litter was present in the substrate. Aquatic roots were formed in the top soil layer when waterlogged. The percentage of aquatic roots increased with increasing amount of litter in the substrate when plants were flooded. It was concluded that the morphological responses of P. australis to litter strongly constrain its ability to maintain itself in deep water when the substrate contains litter. This might one of the explanations for the disappearance of P. australis along the waterward side of littoral zones.     

Effect of submergence on translocation,starch content and amylolytic activity in deep-water rice

Submergence induces rapid internodal elongation in deep-water rice (Oryza sativa L. cv. Habiganj Aman II). We investigated the metabolic activities which help to support such fast growth. Three days of submergence in water under continuous light led to the mobilization of 65% of the starch from those regions of rice internodes which had been formed prior to submergence. Disappearance of starch was accompanied by a 70-fold enhancement of amylolytic activity. Similar increases in amylolytic activity were detected in response to ethylene and gibberellic acid. Submergence also caused a 26-fold increase in the translocation of newly synthesized photosynthetic assimilates from the leaves to the internodes and younger regions of the culms. These physiological processes are likely to provide the metabolic energy required for internodal elongation in response to submergence.Abbreviation GA₃ gibberellic acid     

Physiological and growth responses of Centaurea maculosa (Asteraceae) to root herbivory under varying levels of interspecific plant competition and soil nitrogen availability

Summary Centaurea maculosa seedlings were grown in pots to study the effects of root herbivory by Agapeta zoegana L. (Lep.: Cochylidae) and Cyphocleonus achates Fahr. (Col.: Curculionidae), grass competition and nitrogen shortage (each present or absent), using a full factorial design. The aims of the study were to analyse the impact of root herbivory on plant growth, resource allocation and physiological processes, and to test if these plant responses to herbivory were influenced by plant competition and nitrogen availability. The two root herbivores differed markedly in their impact on plant growth. While feeding by the moth A. zoegana in the root cortex had no effect on shoot and root mass, feeding by the weevil C. achates in the central vascular tissue greatly reduced shoot mass, but not root mass, leading to a reduced shoot/root ratio. The absence of significant effects of the two herbivores on root biomass, despite considerable consumption, indicates that compensatory root growth occurred. Competition with grass affected plant growth more than herbivory and nutrient status, resulting in reduced shoot and root growth, and number of leaves. Nitrogen shortage did not affect plant growth directly but greatly influenced the compensatory capacity of Centaurea maculosa to root herbivory. Under high nitrogen conditions, shoot biomass of plants infested by the weevil was reduced by 30% compared with uninfested plants. However, under poor nitrogen conditions a 63% reduction was observed compared with corresponding controls. Root herbivory was the most important stress factor affecting plant physiology. Besides a relative increase in biomass allocation to the roots, infested plants also showed a significant increase in nitrogen concentration in the roots and a concomitant reduction in leaf nitrogen concentration, reflecting a redirection of the nitrogen to the stronger sink. The level of fructans was greatly reduced in the roots after herbivore feeding. This is thought to be a consequence of their mobilisation to support compensatory root growth. A preliminary model linking the effects of these root herbivores to the physiological processes of C. maculosa is presented.