Evolutionarily stable seasonal timing for insects with competition for renewable resource

Summary I study the evolutionarily stable seasonal patterns of hatching and pupation for herbivorous insects that engage in exploitative competition for a renewable resource. A longer larval feeding period enhances female fecundity, but also causes a higher mortality by predation and parasitism. Previously, it was shown that the evolutionarily stable population exhibits asynchronous starting and ending of the larval feeding period in a model in which larval growth rate decreases with the total larval biomass in the population due presumably to interference competition. Here I study the case in which resource availability changes not only with environmental seasonality but with the depletion by the feeding of larvae. I find that if the impact of the herbivory is strong, both hatching and pupation should occur asynchronously in the evolutionarily stable population. And if the favourable season for the host plant is short the ESS population may include synchronous timing of pupation. If the timing of hatching and pupation occurs asynchronously, in the first day of each interval some fraction of the population hatch or pupate, respectively and the rest do so gradually over the interval. In addition, if the environmental variable changes as a symmetric function of time, the length of the period in which hatching occurs tends to be much shorter than the period in which pupation occurs.     

Stable isotopic niche predicts fitness of prey in a wolf–deer system

Interindividual variation in niche presents a potentially central object on which natural selection can act. This may have important evolutionary implications because habitat use governs a suite of selective forces encountered by foragers. In a free-living native black-tailed deer, Odocoileus hemionus , population from coastal British Columbia, we used stable isotope analysis to identify individual variation in foraging niche and investigated its relationship to fitness. Using an intragenerational comparison of surviving and nonsurviving O. hemionus over 2 years of predation by wolves, Canis lupus, we detected resource-specific fitness. Individuals with isotopic signatures that suggested they foraged primarily in cedar ( Thuja plicata )-dominated and low-elevation hemlock ( Tsuga heterophylla )-dominated forest stands were more likely to be killed by C. lupus . High-quality forage in T. plicata stands, as indexed by protein content, may be involved in maintaining this foraging phenotype. Moreover, nonsurvivors diverged more than survivors from median isotopic signatures, suggesting selection against foraging specialization. Stable isotope analysis provides a novel opportunity to integrate ecological and selective landscapes in order to identify underlying ecological mechanisms of selection and provide insight into the maintenance of variability.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 125–137.     

A stable carbon isotope study of dissolved inorganic carbon cycling in a softwater lake

The dissolved inorganic carbon (DIC) cycle in a softwater lake was studied using natural variations of the stable isotopes of carbon,¹²C and¹³C. During summer stratification there was a progressive decrease in epilimnion DIC concentration with a concomitant increase in ¹³C_DIC), due to preferential uptake of¹²C by phytoplankton and a change in the dominant CO₂ source from inflow andin situ oxidation to invasion from the atmosphere. There was an increase in hypolimnion DIC concentration throughout summer with a concomitant general decrease in ¹³C_DIC from oxidation of the isotopically light particulate organic carbon that sank down through the thermocline from the epilimnion.Mass balance calculations of DI¹²C and DI¹³C in the epilimnion for the summer (June 23–September 25) yield a mean rate of net conversion of DIC to organic carbon (C_org) of 430 ± 150 moles d^-1 (6.5 ± 1.8 m moles m^-2 d^-1. Net CO₂ invasion from the atmosphere was 420 ± 120 moles d^-1 (6.2 ± 1.8 m moles m^-2 d^-1) with an exchange coefficient of 0.6 ± 0.3m d^-1. These results imply that at least for the summer months the phytoplankton obtained about 90% of their carbon from atmosphere CO₂. About 50% of CO₂ invasion and conversion to C_org for the summer occurred during a two week interval in mid-summer.DIC concentration increased in the hypolimnion at a rate of 350 ± 70 moles DIC d^-1 during summer stratification. The amount of DIC added to the hypolimnion was equivalent to 75 ± 20% of net conversion of DIC to C_org in the euphotic zone over spring and summer implying rapid degradation of POC in the hypolimnion. The ¹³C of DIC added to the deep water (-22.) was too heavy to have been derived from oxidation of particulate organic carbon alone. About 20% of the added DIC must have diffused from hypolimnetic sediments where relatively heavy CO₂ (-7) was produced by a combination of POC oxidation and as a by-product of methanogenesis.     

Copper absorption in young men fed adequate and low zinc diets

Copper absorption was measured at two levels of dietary zinc in six healthy young men who were confined to a metabolic unit for a 75 d study of zinc utilization. A diet of conventional foods was fed, providing either 16.5 or 5.5 mg zinc and 1.3 mg copper daily. Copper absorption was determined by feeding⁶⁵Cu, a stable isotope of copper, once during the 16.5 mg Zn diet and near the beginning and end of the 5.5 mg Zn diet. Apparent copper absorption averaged 48.1% when the 16.5 mg Zn diet was fed. This was significantly higher than the averages of 37.2 and 38.5% when the 5.5 mg Zn diet was fed. Absorption also differed significantly among subjects. Fecal copper did not differ between diets or among subjects. All subjects were in positive copper balance at both levels of dietary zinc. These results suggest that a dietary zinc intake slightly above the Recommended Dietary Allowance of 15 mg/d does not increase fecal copper loss and does not interfere with copper absorption.     

In situ response to vinka alkaloids by microtubules in cultured post-implanted mouse embryos

The response of microtubules to treatment with vinca alkaloids was investigated in vivo and in situ in the embryonic nervous system of mice. For this purpose we used rotatory cultures of post-implanted embryos in a serum medium containing the alkaloid combined with immunofluorescence using a tubulin-specific polyclonal antibody on high molecular weight polyethylene glycol embedded semithin sections. In mitotic cells, kinetochore microtubules were seen to be more resistant to the action of vinca alkaloids than interpolar microtubules. Increasing drug concentrations induced an increasing rate of mitosis together with an increasing rate of disassembly of the cytoplasmic microtubule complex, suggesting a probable relation between these events. In bipolar neuroepithelial cells at interphase, a small pool of microtubules was resistant to the vinca alkaloids. These microtubules were located near the centriolar apparatus associated with the primary cilium; they were short, curly and bent. Disruption of the cytoplasmic microtubule complex did not alter the shape of the bipolar neuroepithelial cells. In the axonal profiles, a drug-stable pool of microtubules were not disrupted by the alkaloids and were also short. They seem to act as microtubule organizing centres. These observations suggest vinca alkaloids seem to act in vivo much more by inducing, at a given concentration, the disruption of a particular group of microtubules without altering the others. The fact that these drugs affect the number, but not the length, of the microtubules raises the hypothesis that these drugs act on microtubules by a mechanism similar to that described as "dynamic instability".     

Seasonal water uptake and movement in root systems of Australian phraeatophytic plants of dimorphic root morphology: a stable isotope investigation

A natural abundance hydrogen stable isotope technique was used to study seasonal changes in source water utilization and water movement in the xylem of dimorphic root systems and stem bases of several woody shrubs or trees in mediterranean-type ecosystems of south Western Australia. Samples collected from the native treeBanksia prionotes over 18 months indicated that shallow lateral roots and deeply penetrating tap (sinker) roots obtained water of different origins over the course of a winter-wet/summer-dry annual cycle. During the wet season lateral roots acquired water mostly by uptake of recent precipitation (rain water) contained within the upper soil layers, and tap roots derived water from the underlying water table. The shoot obtained a mixture of these two water sources. As the dry season approached dependence on recent rain water decreased while that on ground water increased. In high summer, shallow lateral roots remained well-hydrated and shoots well supplied with ground water taken up by the tap root. This enabled plants to continue transpiration and carbon assimilation and thus complete their seasonal extension growth during the long (4–6 month) dry season. Parallel studies of other native species and two plantation-grown species ofEucalyptus all demonstrated behavior similar to that ofB. prionotes. ForB. prionotes, there was a strong negative correlation between the percentage of water in the stem base of a plant which was derived from the tap root (ground water) and the amount of precipitation which fell at the site. These data suggested that during the dry season plants derive the majority of the water they use from deeper sources while in the wet season most of the water they use is derived from shallower sources supplied by lateral roots in the upper soil layers. The data collected in this study supported the notion that the dimorphic rooting habit can be advantageous for large woody species of floristically-rich, open, woodlands and heathlands where the acquisition of seasonally limited water is at a premium.     

15N natural abundance in forest and pasture soils of the Brazilian Amazon Basin

The natural abundance of ¹⁵N was examined in soil profiles from forests and pastures of the Brazilian Amazon Basin to compare tropical forests on a variety of soil types and to investigate changes in the sources of nitrogen to soils following deforestation for cattle ranching. Six sites in the state of Rondônia, two sites in Pará and one in Amazonas were studied. All sites except one were chronosequences and contained native forest and one or more pastures ranging from 2 to 27 years old. Forest soil ¹⁵N values to a depth of 1 m ranged from 8 to 23 and were higher than values typically found in temperate forests. A general pattern of increasing ¹⁵N values with depth near the soil surface was broadly similar to patterns in other forests but a decrease in ¹⁵N values in many forest profiles between 20 and 40 cm suggests that illuviation of ¹⁵N-depleted nitrate may influence total soil ¹⁵N values in deeper soil where total N concentrations are low. In four chronosequences in Rondônia, the ¹⁵N values of surface soil from pastures were lower than in the original forest and ¹⁵N values were increasingly depleted in older pastures. Inputs of atmospheric N by dinitrogen fixation could be an important N source in these pastures. Other pastures in Amazonas and Pará and Rondônia showed no consistent change from forest values. The extent of fractionation that leads to ¹⁵N enrichment in soils was broadly similar over a wide range of soil textures and indicated that similar processes control N fractionation and loss under tropical forest over a broad geographic region. Forest ¹⁵N profiles were consistent with conceptual models that explain enrichment of soil ¹⁵N values by selective loss of ¹⁴N during nitrification and denitrification.     

A pyrenoid-based carbon-concentrating mechanism is present in terrestrial bryophytes of the class Anthocerotae

It has been widely accepted that carbon assimilation in bryophytes is exclusively based on the conventional C₃ photosynthetic pathway. The occurrence of biochemical CO₂-concentrating mechanisms (C₄ or Crassulacean acid metabolism), which have developed in plants in the last 20–100 million years, has been discounted for bryophytes from studies of the carbon isotope composition (¹³C) of organic material. In contrast cyanobacteria and many algae show active accumulation of dissolved inorganic carbon via biophysical CO₂-concentrating mechanisms which are also found in the photobiont partners in certain lichens. The presence of a pyrenoid, a granular particle within the chloroplast, has been linked with CO₂-concentrating mechanism activity in green algae and lichens and we now show that such a mechanism is categorically associated with the occurrence of a pyrenoid in bryophytes belonging to the class of Anthocerotae. These observations have significant evolutionary implications for the development of terrestrial photosynthesis during the colonisation of the land, raising the intriguing question of why the pyrenoid-based CO₂-concentrating mechanism did not persist in the terrestrial environment.Abbreviations and Symbols CCM carbon-concentrating mechanism - DIG dissolved inorganic carbon (CO₂+HCO ₃ ^- +CO ₂ ^- ) - DW dry weight - K_0.5 external concentration of CO₂ at which half-maximal rates of CO₂ assimilation are reached - Rubisco ribulose-l,5-bisphosphate carboxylase-oxygenase - carbon isotope discrimination (%) - ¹³C carbon isotope ratio (%) This work was supported by the Natural Environment Research Council (GR3/8813) and the Leverhulme Trust. We thank Prof. A. Roy Perry (National Museum of Wales, Cardiff), Dr. B. Coppins and Mr. D. Long (Royal Botanic Garden Edinburgh) for access to herbarium specimens and Mr. M. Fletcher for providing living bryophytes.     

Variations in stream water uptake by Eucalyptus camaldulensis with differing access to stream water

The stable isotopes ²H and ¹⁸O were used to determine the water sources of Eucalyptus camaldulensis at three sites with varying exposure to stream water, all underlain by moderately saline groundwater. Water uptake patterns were a function of the long-term availability of surface water. Trees with permanent access to a stream used some stream water at all times. However, water from soils or the water table commonly made up 50% of these trees' water. Trees beside an ephemeral stream had access to the stream 40–50% of the time (depending on the level of the stream). No more than 30% of the water they used was stream water when it was available. However, stream water use did not vary greatly whether the trees had access to the stream for 2 weeks or 10 months prior to sampling. Trees at the third site only had access to surface water during a flood. These trees did not change their uptake patterns during 2 months inundation compared with dry times, so were not utilising the low-salinity flood water. Pre-dawn leaf water potentials and leaf ¹³C measurements showed that the trees with permanent access to the stream experienced lower water stress and had lower water use efficiencies than trees at the least frequently flooded site. The trees beside the ephemeral stream appeared to change their water use efficiency in response to the availability of surface water; it was similar to the perennial-stream trees when stream water was available and higher at other times. Despite causing water stress, uptake of soil water and groundwater would be advantageous to E. camaldulensis in this semi-arid area, as it would provide the trees with a supply of nutrients and a reliable source of water. E. camaldulensis at the study site may not be as vulnerable to changes in stream flow and water quality as previously thought.