Growth and competition of Cytisus scoparius,an invasive shrub,and Australian native shrubs

English broom (Cytisus scoparius) is an aggressive invasive shrub in native sclerophyll forests of South Australia. We studied its relative growth rate (RGR) and competitive ability in soils from invaded and uninvaded woodlands, in comparison to three native species it commonly displaces:Hakea rostrata, Acacia verniciflua, and A. myrtifolia. Hakea was the slowest growing species throughout the year. Both native species had their highest RGR during spring. The RGR of broom was higher than that of both hakea and acacia in the winter and spring. Despite losing its leaves in the summer, the RGR of broom through the year was higher than that of either of the native species. Soil from the invaded stands had higher organic C, N and soluble P than that from uninvaded sites. Broom and acacia grew better in the higher nutrient soil than in the lower nutrient soil. Competition did not decrease the final biomass of any of the species in low nutrient soil. In the higher nutrient soil the biomass of broom was reduced by competition with acacia, but not by competition with hakea. Competition by broom reduced the biomass of hakea but not that of acacia. Broom's earlier and higher RGR, high competitiveness in nutrient rich soils, and probably its ability to change nutrient availability could be important contributors to the mechanisms by which it invades native woodlands.     

Efficient clonal propagation method for Decalepis hamiltonii, an endangered shrub, under the influence of phloroglucinol

A highly efficient two stage protocol was developed for induction of multiple shoots from single node in vitro shoot tip explants of Decalepis hamiltonii. It was found that phloroglucinol (PG) had synergistic effect on shoot multiplication when added with N6-benzyladenine and gibberellic acid. This protocol uses PG for both multiple shoot induction from nodal explants, elongation of primary shoots and initiation of adventitious shoot formation from primary shoots, which was more in presence of triacontanol (TRIA). Maximum number of shoots per culture was observed on the medium containing N6-benzyladenine (1.1 microM; BA), GA3 (5.8 microM) and PG (800 microM). Sub-culturing of the shoots onto MS medium containing optimum concentration of BA (5.6 microM), PG (200 microM) and TRIA (0.011 microM) produced elongated shoots along with secondary shoot formation. The long shoots were rooted on alpha-naphthalene acetic acid (5.38 microM; NAA) and PG (400 microM) containing medium. The rooted plantlets were hardened and their field survival rate was 80-90%.     

Population density and module demography in Trapa natans (Trapaceae), an annual,clonal aquatic macrophyte

The effects of population density on module demography were studied in Trapa natans L., an annual aquatic macrophyte capable of extensive clonal propagation. At low density, the floating plants produced ten times as many ramets (clonal offshoots) as those at high density. Module mortality occurred at three levels: leaf, ramet (shoot), and genet (genetic individual). There was 100-fold variation in the size of nuts containing germinable seeds. In early summer there was a highly significant linear relationship between dry mass of nuts and the total mass of ramets that each had produced. In early summer most (73–83%) of the variation in total plant biomass was attributable to variation in initial nut size. However the significance of initial nut size was diminished later in the season. The great success of the exotic weed T. natans at colonizing and monopolizing an aquatic habitat is a function of its highly productive clonal growth response to low-density conditions, combined with greater proportional allocation of biomass to reproductive structures, resulting in greatly increased nut production at low initial density. The species appears able to develop and maintain a population at extremely high density: plant buoyancy and the production of large, well-protected nuts allow rapid early growth from the sediment each year and early pre-emption of the water surface.     

Onset rivalry: brief presentation isolates an early independent phase of perceptual competition

When the left and right eyes are simultaneously presented with different images, observers typically report exclusive awareness of only one image. This phenomenon is termed binocular rivalry, reflecting the fact that the dominant image alternates every few seconds in a cycle of perceptual competition that continues indefinitely. Despite the apparent continuity in perceptual switching, we now demonstrate that the initial "onset" period is fundamentally different to all subsequent rivalry epochs. Using brief intermittent presentations, rivalry dominance shows strong biases such that the same target is perceived with each successive stimulus onset. These biases remain consistent within any given location, but vary across the visual field in a distribution that is stable over multiple weeks but highly idiosyncratic across observers. If the presentation exceeds approximately 1sec at any location, however, the very different and much more balanced alternations of sustained binocular rivalry become apparent. These powerful onset biases are observed with brief intermittent presentations at a single location or with continual smooth motion of the targets. Periods of adaptation to one of the rivaling targets induced local switches in dominance to the non-adapted target. However, these effects were generally limited to the spatial site of adaptation and had less influence over each subsequent cycle of the target. We conclude that onset rivalry is independent of sustained rivalry and cannot be explained by local regions of monocular dominance or memory of past perceptual history, but rather reflects low-level, spatially localized factors that are stable over periods of weeks. These findings suggest that brief presentation paradigms are inappropriate for their current use in studies of the mechanisms underlying sustained rivalry. However, brief presentations are ideal for investigating early stages of perceptual competition.     

Patch structure and ramet demography of the clonal tree, Asimina triloba, under gap and closed-canopy

Clonal understory trees develop into patches of interconnected and genetically identical ramets that have the potential to persist for decades or centuries. These patches develop beneath forest canopies that are structurally heterogeneous in space and time. Canopy heterogeneity, in turn, is responsible for the highly variable understory light environment that is typically associated with deciduous forests. We investigated what aspects of patch structure (density, size structure, and reproductive frequency of ramets) of the clonal understory tree, Asimina triloba, were correlated with forest canopy conditions. Specifically, we compared A. triloba patches located beneath closed canopies and canopy gaps. We also conducted a three-year demographic study of individual ramets within patches distributed across a light gradient. The closed canopy-gap comparison demonstrated that the patches of A. triloba had a higher frequency of large and flowering ramets in gaps compared to closed-canopy stands, but total ramet density was lower in gaps than in closed canopy stands. In the demographic study, individual ramet growth was positively correlated with light availability, although the pattern was not consistent for all years. Neither ramet recruitment nor mortality was correlated with light conditions. Our results indicate that the structure of A. triloba patches was influenced by canopy condition, but does not necessarily depend on the responses of ramets to current light conditions. The lack of differences in ramet recruitment and mortality under varying canopy conditions is likely to be a primary reason for the long-term expansion and persistence of the patches. The primary benefit of a positive growth response to increasing light is the transition of relatively small ramets into flowering ramets within a short period of time.     

Fine-scale clonal structure and diversity within patches of a clone-forming dioecious shrub, Ilex leucoclada (Aquifoliaceae)

BACKGROUND AND AIMS: The mode of reproduction (sexual vs. asexual) is likely to have important effects on genetic variation and its spatial distribution within plant populations. An investigation was undertaken of fine-scale clonal structure and diversity within patches of Ilex leucoclada (a clone-forming dioecious shrub). METHODS: Six patches were selected in a 1-ha plot previously established in an old-growth beech forest. Two of the selected patches were composed predominantly of stems with male flowers (male patch), and two contained stems with predominantly female flowers (female patch). The remaining two patches contained stems with male flowers and stems with female flowers in more or less equal proportions (mixed patch). Different genets were distinguished using random amplified polymorphic DNA (RAPD) markers. KEY RESULTS: One hundred and fifty-six genets with different RAPD phenotypes were identified among 1928 stems from the six patches. Among the six patches, the male patches had the lowest clonal diversity, and the mixed patches had the highest. Distribution maps of the genets showed that they extended downhill, reflecting natural layering that occurred when stems were pressed to the ground by heavy snow. In every patch, there were a few large genets with many stems and many small genets with a few stems. CONCLUSION: The differences in clonal diversity among patches may be due to differences in seedling recruitment frequencies. The skewed distribution of genet size (defined as the number of stems per genet) within patches may be due to differences in the timing of germination, or age (with early-establishing genets having clear advantages for acquiring resources) and/or intraspecific competition.     

The effect of water and nitrogen amendments on photosynthesis,leaf demography,and resource-use efficiency in Larrea tridentata,a desert evergreen shrub

Summary In the Chihuahuan Desert of southern New Mexico, both water and nitrogen limit the primary productivity of Larrea tridentata, a xerophytic evergreen shrub. Net photosynthesis was positively correlated to leaf N, but only in plants that received supplemental water. Nutrient-use efficiency, defined as photosynthetic carbon gain per unit N invested in leaf tissue, declined with increasing leaf N. However, water-use efficiency, defined as the ratio of photosynthesis to transpiration, increased with increasing leaf N, and thus these two measures of resource-use efficiency were inversely correlated. Resorption efficiency was not significantly altered over the nutrient gradient, nor was it affected by irrigation treatments. Leaf longevity decreased significantly with fertilization although the absolute magnitude of this decrease was fairly small, in part due to a large background of insect-induced mortality. Age-specific gas exchange measurements support the hypothesis that leaf aging represents a redistribution of resources, rather than actual deterioration or declining resource-use efficiency.     

The demography of a clonal herbaceous perennial plant, the longleaved sundew Drosera intermedia, in different heathland habitats

Eight populations of longleaved sundew Drosera intermedia Hayne ramets were studied in three hydrologically and phytosociologically differing heathland habitats over a 4 yr period a pool edge, an old path through the wet heath, a seepage area At the extreme sites, with high levels of water table fluctuations (pool edge and dry path), both survival and recruitment were low, resulting in small populations The pool edge populations went extinct At the less extreme sites (wet paths and seepage area), populations were stable, but ramet and genet turnover was higher at the path sites than in the seepage area Demographic variation between populations was explained by differences in adult survival and sexual recruitment (asexual recruitment did not differ clearly), which, in turn, were related to specific biotic and abiotic factors Likely adult mortality causes were summer drought (all habitats), winter frost (path habitat), cover with a crust of iron bacteria after heavy summer rainfall (seepage area) and shading by Sphagnum and algae (pool edge) Low seedhng recruitment was caused by a low germination succes (pool, seepage area) and seedling survival (seepage area, dry path)     

Two decades of demography reveals that seed and seedling transitions limit population persistence in a translocated shrub

Background and Aims

Olearia flocktoniae is an endangered shrub that was passively translocated from its natural ecosystem, where it has since gone extinct. This study aimed to determine sensitivities vital to populations persisting in human-created areas.

Methods

Population colonization, longevity and extinction were investigated over 20 years using 133 populations. Seed-bank longevity was determined from germination trials of seeds exhumed from extinct and extant sites via a 10-year glasshouse trial and by in situ sowing experiments. From 27 populations, 98 cohorts were followed and matrix models of transitions from seeds to adults were used to evaluate the intrinsic rate of population growth against disturbance histories. Ten populations (38 cohorts) with different disturbance histories were used to evaluate sensitivities in vital rates.

Key Results

Most populations had few individuals (∼30) and were transient (<5 years above ground). The intrinsic population growth rate was rarely >1 and all but two populations were extinct at year 20. Seeds were short-lived in situ. Although >1000 seeds per plant were produced annually in most populations, sensitivity analysis showed that the transition to the seed bank and the transition from the seed bank to seedlings are key vulnerabilities in the life-cycle.

Conclusions

Seedling establishment is promoted by recent disturbance. Increasing the number of disturbance events in populations, even severe disturbances that almost extirpate populations, significantly increases longer-term population persistence. Only populations that were disturbed annually survived the full 20 years of the study. The results show that translocated populations of O. flocktoniae will fail to persist without active management.     

Disturbance,competition and the maintenance of clonal diversity in Daphnia pulex

Laboratory microcosm experiments tested the intermediate disturbance hypothesis, which states that the highest level of diversity (e.g. species diversity) will be maintained at intermediate scales of disturbance. The effects of disturbance on the maintenance of clonal diversity and on competitive interactions among clones of the obligately parthenogenetic freshwater cladoceran, Daphnia pulex were examined. No significant effect of disturbance size (i.e. dilution volume) on clonal diversity was noted. However, frequency of disturbance had a pronounced effect on clonal diversity, with the highest clonal diversity maintained at low to intermediate disturbance frequencies. Competitive hierarchies among clones were often invariant within a given experiment. Generally, one or two clones dominated, with several less abundant clones persisting throughout an experiment. Results suggest that low to intermediate disturbances could be important in the maintenance of genetic variation in natural populations (i.e. through pre-emption of competitive exclusion between genotypes). This could have a direct bearing on the maintenance of both intra- and interspecific diversity.     

Coexisting congeners: demography, competition, and interactions with cardenolides for two milkweed-feeding aphids

Explaining the coexistence of closely related species sharing a single resource has been a long-standing challenge in ecology. Here we report on studies comparing the aphids Aphis nerii and A. asclepiadis that feed sympatrically on the milkweed Asclepias syriaca in northeastern North America. We sought to identify tradeoffs among species' attributes that might promote coexistence, but in most instances A. nerii was superior to A. asclepiadis . Aphis nerii was 84% more fecund, fed upon 880% more phloem sap, and was affected 70% less by intraspecific competition as compared to A. asclepiadis . In interspecific competition, A. nerii reduced A. asclepiadis abundance by 77%, whereas A. asclepiadis did not affect A. nerii . In dispersal trials, 10% of winged A. nerii but only 1% of A. asclepiadis successfully moved from non-host plants to A. syriaca . We also investigated whether there were differences in aphid interactions with milkweed cardenolides. Jasmonic acid increased milkweed cardenolides by 33%, a realistic amount similar to that induced by several leaf-chewing herbivores. Nevertheless, jasmonate-induced cardenolides failed to affect aphid performance and aphid feeding had no effect on milkweed cardenolide concentration. Yet cardenolides were important for aphid resistance to predators; A. nerii sequestered 25% more cardenolides and was preyed upon 50% less than A. asclepiadis . Interactions with cardenolides thus again favored A. nerii over A. asclepiadis . Given that A. nerii and A. asclepiadis are decidedly not equivalent in their demography, competitive ability, defense and dispersal, our results strongly refute the notion that neutral processes can explain coexistence of these aphids. Based on field observations, we propose two tradeoffs – timing of milkweed colonization and relationships with ants – as putative mechanisms for the coexistence of these congeners.     

Responses to harvesting intensity in a clonal dwarf shrub,the bilberry (Vaccinium myrtillus L.)

The effects of herbivory were simulated on stands of bilberry (Vaccinium myrtillus L.) in a boreal Empetrum-Myrtillus type forest. Five harvesting intensities were used (0% (A), 25% (B), 50% (C), 100% (D) of the bilberry ramets or all the ramets of all species (E)). Density and biomass of the stands, and growth of the ramets were monitored for five growth seasons, from 1986 to 1990. After damage new ramets emerged rapidly from dormant buds at the base of removed ramets. Between 70 and 97% of the density relative to the control level was regained by the final harvest. However, only between 11 and 64% of the biomass relative to the control level was recovered. Clipping reduced the branch growth, both in the new ramets and in the new parts of the old, unclipped ramets. Severe treatments (D and E) decreased the growth more than did light harvesting (B and C). Survival and fecundity of the ramets were not affected. The result therefore suggested that the bilberry is not able to recover totally from severe herbivory damage between the years of peak rodent population that shows a 3–4 year cycle. Nevertheless, extreme grazing pressure is rare in boreal ecosystems as alternative food is available for herbivores, and predators also limit the herbivore population. Hence the bilberry exhibits moderate tolerance of the usual level of herbivory damage.     

Comparative responses of a non-N-fixing shrub and an actinorhizal N-fixing shrub to N fertilization

Background and aims

Variations in responses to soil N between a non-N-fixing shrub, Baccharis halimifolia L., and a N-fixing shrub, Morella cerifera (L.) Small, were tested over 12 weeks to determine whether N availability is the sole cause of persistent dominance of M. cerifera on barrier islands.

Methods

Plants were supplied increasing levels of soil N up to 200 mg kg^?1. Measurements included gas exchange and chlorophyll fluorescence parameters across treatments, species, and time. Tissues were analyzed for differences in biomass and nutrients.

Results

Baccharis halimifolia had reduced physiological responses across all treatment levels, but M. cerifera had comparatively few variations. Across all treatments B. halimifolia photosynthesis and stomatal conductance were reduced by 62 and 76 %, respectively,by week 12. Increasing foliar δ¹⁵N values across treatments for M. cerifera indicated a shift from utilizing fixed N to available soil N. Biomass was highest at 200 mg kg^?1 N for both species. Baccharis halimifolia showed indications of stress response and resource limitation based on physiological responses, nutrient contents, and isotope effects.

Conclusions

Baccharis halimifolia showed signs of co-limitation of both N and P whereas M. cerifera was limited by neither, suggesting that dominance of M. cerifera is only partially explained by actinorhizal symbiosis and N availability.     

T cell clonal conditioning: a phase occurring early after antigen presentation but before clonal expansion is impacted by Toll-like receptor stimulation

After in vivo immunization, Ag-specific T cells disappear from circulation and become sequestered in lymphoid tissue where they encounter Ag presented by dendritic cells. In the same site and just after Ag presentation, they "disappear" a second time and we investigated this process. Using a mouse model of T cell deletion (without Toll-like receptor (TLR) stimulation) vs survival (with TLR stimulation), Ag-specific T cells indeed became undetectable by flow cytometry, however were readily detected by immunohistochemistry. Thus, whether or not the activated T cells were destined to delete or survive, they were difficult to extract from lymphoid tissue and did not disappear but in fact were abundantly present. Nevertheless, profound differences were observed during this time period when tolerizing conditions were compared with immunizing conditions. TLR stimulation induced an increase in CD25 expression, acquisition of surface MHC class II, and abnormally high increases in forward and side scatter of the peptide-specific T cells. Using a modified adoptive transfer approach, we demonstrated by flow cytometry that in the presence of TLR stimulation the Ag-specific T cells were tightly coupled to dendritic cells, explaining the unusual increases in size and granularity. Ultimately, these events induced the specific T cells to differentiate into memory cells. We postulate that this is a stage where T cells are either conditioned to survive or to delete depending upon the activation status of the innate immune system.     

Translocation and early post-release demography of endangered Laysan teal

In an attempt to reduce the high extinction risk inherent to small island populations, we translocated wild Laysan teal Anas laysanensis to a portion of its presumed prehistoric range. Most avian translocations lack the strategic post-release monitoring needed to assess early population establishment or failure. Therefore, we monitored the survival and reproduction of all founders, and their first-generation offspring using radio telemetry for 2 years after the first release. Forty-two Laysan teal were sourced directly from the only extant population on Laysan Island and transported 2 days by ship to Midway Atoll. All birds survived the translocation with nutritional and veterinary support, and spent between 4 and 14 days in captivity. Post-release survival of 42 founders was 0.857 (95% CI 0.86–0.99) during 2004–2006 or annualized 0.92 (95% CI 0.83–0.98). Seventeen of 18 founding hens attempted nesting in the first two breeding seasons. Fledgling success was 0.57 (95% CI 0.55–0.60) in 2005 and 0.63 (95% CI 0.62–0.64) in 2006. The effective founding female population ( N _e) was 13. We applied these initial demographic rates to model population growth. The nascent population size increased to >100 after only 2 years post-release ( λ =1.73). If this growth rate continues, the size of the Midway population could surpass the source population before 2010.     

Biomass allocation in a clonal vine: Effects of intraspecific competition and nutrient availability

Biomass allocation to roots, rhizomes, runners and climbing stems (i.e. twining axis and attached leaves) was studied inCalystegia sepium L., a clonal vine. In an experiment which took 2 months, nutrient availability (low and high) and intraspecific competition (none, shoot root and both shoot and root) were manipulated. Under low nutrients the highest biomass of climbing stems was found in plants with shoot competition; the lowest was found in plants with both shoot and root competition. Total biomass under high nutrients was also greatest in plants with shoot competition. Thus, plants benefited from climbing up a shared stake rather than separate stakes. Larger plants allocated a higher proportion of biomass to runners in the nutrient-poor environment than in the nutrient-rich environment. This behaviour may increase the chance of finding nutrient-rich patches in the neighbourhood of the mother plant in a heterogeneous environment.     

Effects of intraspecific competition on size variation and reproductive allocation in a clonal plant

Clonal plants grow in diameter rather than height, and therefore competition among genets is likely to be symmetric and to result in smaller variation in size of genets than in non-clonal plants. Moreover, clonal plants can reproduce both sexually and vegetatively. We studied the effects of density on the size of rosettes and of clones, variation in the size of rosettes and of clones, and allocation to sexual and vegetative reproduction in the clonal herb Ranunculus reptans . We grew plants from an artificial population of R. reptans in 32 trays at two densities. After four months, differences in density were still apparent, although clones in the low-density treatment had on average 155% more rosettes and 227% more rooted rosettes than clones in the high-density treatment. The coefficient of variation of these measures of clone size was 15% and 83% higher, respectively, in the low-density treatment. This indicates that intraspecific competition among clones of R. reptans is symmetric and increases the effective population size. Rooted rosettes were larger and varied more in size in the low-density treatment. The relative allocation of the populations to sexual and to vegetative reproduction was 19% and 13% higher, respectively, in the high-density treatment. Moreover, seeds produced in the high-density treatment had a 24% higher mass and a 7% higher germination percentage. This suggests that with increasing density, allocation to sexual reproduction increases more than allocation to vegetative reproduction in R. reptans , which corresponds to the response of some other species with a spreading growth form but not of species with a compact growth form. We conclude that intraspecific competition is an important factor in the life-history evolution of R. reptans because intraspecific competition affects its clonal life-history traits and may affect evolutionary processes such as genetic drift and selection through its effect on the effective population size.     

Leaf demography of the rainforest shrub Wilkiea macrophylla and its implications for the ecology of foliicolous lichens

A population of leaves of the rainforest shrub Wilkiea macrophylla was studied for a period of 3 years. Survival curves for three subsamples were constructed. The estimated proportion of leaves initiated during the study that survived for the length of the study period (1150 days) was 79.7% and, by extrapolating in time, their mean half life was estimated to be 6.8 years. Those leaves present at the inception of the programmes but without lichen cover at that time, showed 80.1% survival over the subsequent 1150 days, whereas only 55.2% of those with lichen cover survived for 1150 days. Leaf growth and fall was concentrated in the period September-December, suggesting cool temperate affinities. Leaves of W. macrophylla are so long-lived that it cannot be assumed that foliicolous lichens are ruderals: only comparative studies will establish their nature.     

Spatial heterogeneity in light supply affects intraspecific competition of a stoloniferous clonal plant

Spatial heterogeneity in light supply is common in nature. Many studies have examined the effects of heterogeneous light supply on growth, morphology, physiology and biomass allocation of clonal plants, but few have tested those effects on intraspecific competition. In a greenhouse experiment, we grew one (no competition) or nine ramets (with intraspecific competition) of a stoloniferous clonal plant, Duchesnea indica, in three homogeneous light conditions (high, medium and low light intensity) and two heterogeneous ones differing in patch size (large and small patch treatments). The total light in the two heterogeneous treatments was the same as that in the homogeneous medium light treatment. Both decreasing light intensity and intraspecific competition significantly decreased the growth (biomass, number of ramets and total stolon length) of D. indica. As compared with the homogeneous medium light treatment, the large patch treatment significantly increased the growth of D. indica without intraspecific competition. However, the growth of D. indica with competition did not differ among the homogeneous medium light, the large and the small patch treatments. Consequently, light heterogeneity significantly increased intraspecific competition intensity, as measured by the decreased log response ratio. These results suggest that spatial heterogeneity in light supply can alter intraspecific interactions of clonal plants.