Adaptive advantages of patterns of growth and asexual reproduction of the sea anemone Metridium senile (L.) in intertidal and submerged populations

Individual size, rate of growth, and mode and frequency of asexual reproduction are life-history traits of primary importance for sea anemones. These traits determine sexual reproductive output, affect an individual's probability of survival, and are crucial in adapting an individual to its environmental surroundings. The sea anemone Metridium senile (L.) is highly variable in ecological distribution and life history, including rate of growth, individual size, and rate of asexual reproduction. Gonad size (measured as cross-sectional area of gonadal tissue) increases with body weight, so individuals should grow as large and as rapidly as possible to maximize individual sexual reproductive output. Cessation of growth and small body size in intertidal populations suggest that growth is constrained by genetic or environmental conditions. The growth of intertidal individuals transplanted to harbor-float panels demonstrated that growth limits are imposed by environmental factors, most probably limited food and feeding time and damage from wave exposure (which stimulates fragmentation). Individuals in harbor-float populations, which are continuously immersed, grow much larger, and large individuals comprise a greater proportion of the population than in the intertidal zone. The highest rate of fragmentation observed was on harbor-float panels. Patterns of growth and asexual reproduction provide adaptive advantages for M. senile. For harborfloat individuals, large individual size increases gamete production and may increase feeding efficiency. For intertidal individuals, asexual reproduction allows growth despite individual size constraints and rapid population growth, with specific advantages resulting from clone formation.     

Trade-offs between sexual and asexual reproduction in the genus Mimulus

Summary Six estimates of resource allocation to sexual reproduction (nectar concentration, nectar volume, sugar production, pollinator visitation rates, fruit-set, and seed production) and a single estimate of resource allocation to asexual reproduction (the rate of rooted-branch production) were examined for five species of Mimulus. There were significant interspecific differences for all paramaters. With the exception of nectar concentration, there was 1) a consistent positive correlation among the parameters measuring allocation to sexual reproduction, and 2) a significant negative correlation between parameters measuring allocation to sexual reproduction and the rate of rooted-branch production (asexual reproduction). The results indicate that Mimulus species which produce the highest volumes of nectar 1) receive the most pollinator visits, 2) have the highest fruit-set, 3) produce the most seeds, and 4) produce the fewest rooted branches.     

The conservation of redundancy in genetic systems: effects of sexual and asexual reproduction

The relationship between probability of survival and the number of deleterious mutations in the genome is investigated using three different models of highly redundant systems that interact with a threatening environment. Model one is a system that counters a potentially lethal infection; it has multiple identical components that act in sequence and in parallel. Model two has many different overlapping components that provide three-fold coverage of a large number of vital functions. The third model is based on statistical decision theory: an ideal detector, following an optimum decision strategy, makes crucial decisions in an uncertain world. The probability of a fatal error is reduced by a redundant sampling system, but the chance of error rises as the system is impaired by deleterious mutations. In all three cases the survival profile shows a synergistic pattern in that the probability of survival falls slowly and then more rapidly. This is different than the multiplicative or independent survival profile that is often used in mathematical models. It is suggested that a synergistic profile is a property of redundant systems. Model one is then used to study the conservation of redundancy during sexual and asexual reproduction. A unicellular haploid organism reproducing asexually retains redundancy when the mutation rate is very low (0001 per cell division), but tends to lose high levels of redundancy if the mutation rate is increased (001 to 01 per cell division). If a similar unicellular haploid organism has a sexual phase then redundancy is retained for mutation rates between 0001 and 01 per cell division. The sexual organism outgrows the asexual organism when the above mutation rates apply. If they compete for finite resources the asexual organism will be extinguished. Variants of the sexual organism with increased redundancy will outgrow those with lower levels of redundancy and the sexual process facilitates the evolution of more complex forms. There is a limit to the extent that complexity can be increased by increasing the size of the genome and in asexual organisms this leads to progressive accumulation of mutations with loss of redundancy and eventual extinction. If complexity is increased by using genes in new combinations, the asexual form can reach a stable equilibrium, although it is associated with some loss of redundancy. The sexual form, by comparison, can survive, with retention of redundancy, even if the mutation rate is above one per generation. The conservation and evolution of redundancy, which is essential for complexity, depends on the sexual process of reproduction.     

Latitudinal variation in plant size and relative growth rate in Arabidopsis thaliana

Latitude is an important determinant of local environmental conditions that affect plant growth. Forty ecotypes of Arabidopsis thaliana were selected from a wide range of latitudes (from 16°N to 63°N) to investigate genetic variation in plant size and relative growth rate (RGR) along a latitudinal gradient. Plants were grown in a greenhouse for 31 days, during which period three consecutive harvests were performed. Plants from high latitudes tended to have smaller plant size in terms of seed size, cotyledon width, rosette size, number of rosette leaves, size (leaf area) of the largest leaves, total leaf area, and total dry weight per plant than those from low latitudes. The mean (±SE) RGR across ecotypes was 0.229 (±0.0013) day⁻¹. There was, however, significant ecotypic variation, with RGR being negatively correlated with latitude. The two main components of RGR, leaf area ratio (LAR) and unit leaf rate (ULR), were also correlated with latitude: LAR increased with increasing latitude while ULR decreased with increasing latitude. It was also found that RGR tended to be negatively correlated with LAR, specific leaf area (SLA) and specific root length (SRL) but to be positively correlated with mean area per leaf (MAL) and ULR. The variation in RGR among ecotypes was relatively small compared with that in the other traits. RGR may be a conservative trait, whose variation is constrained by the trade-off between its physiological (i.e. ULR) and morphological (i.e. LAR) components. Received: 2 November 1997 / Accepted: 28 February 1998     

Contrasting patterns of synonymous and nonsynonymous sequence evolution in asexual and sexual freshwater snail lineages

In asexual lineages, both synonymous and nonsynonymous sequence polymorphism may be reduced due to severe founder effects when asexual lineages originate. However, mildly deleterious (nonsynonymous) mutations may accumulate after asexual lineages are formed, because the efficiency of purifying selection is reduced even in the nonrecombining mitochondrial genome. Here we examine patterns of synonymous and nonsynonymous mitochondrial sequence polymorphism in asexual and sexual lineages of the freshwater snail Campeloma. Using clade-specific estimates, we found that synonymous sequence polymorphism was significantly reduced by 75% in asexuals relative to sexuals, whereas nonsynonymous sequence polymorphism did not differ significantly between sexuals and asexuals. Two asexual clades had high negative values for Tajima's D statistic. Coalescent simulations confirmed that various bottleneck scenarios can account for this result. We also used branch-specific estimates of the ratio of amino acid to silent substitutions, K(a)/K(s). Our study revealed that K(a)/K(s) ratios are six times higher in terminal branches of independent asexual lineages compared to sexuals. Coalescent-based reconstruction of gene networks for all sexual and asexual clades indicated that nonsynonymous mutations occurred at a higher frequency in recently derived asexual haplotypes. These findings suggest that patterns of synonymous and nonsynonymous nucleotide polymorphism in asexual snail lineages may be shaped by both severe founder effect and relaxed purifying selection.     

The influence of population density on fission and growth of Holothuria atra in natural mesocosms

Investigation of the population dynamics, asexual reproduction (fission) and growth of holothuroids has been impeded by the difficulty of tagging individuals. We conducted the first tests on the interactions between population density, fission and growth of holothuroids in experimental populations placed in natural mesocosms (microatolls) at One Tree Reef (OTR), Great Barrier Reef. Similarly sized Holothuria atra were translocated to the microatolls in low (LDT) and high (HDT) density treatments. We hypothesised that holothuroids in lower density treatments would have more resources per individual and that this would promote higher frequencies of asexual reproduction and smaller individuals. The seasonal pattern of fission was similar in natural (unmanipulated) and experimental populations, with the maximum number of fission products occurring in winter and spring. The overall density of the LDT (0.19 ind. m^- 2) and HDT (0.59 ind. m^- 2) did not vary over time. This ‘steady state’ suggested that some fission products died and that asexual reproduction compensated for overall mortality and emigration. There was no difference in sediment chlorophyll pigments between treatments indicating that the different densities of H. atra did not affect benthic microalgal biomass. The percentage of fission products was greater in the LDT than the HDT but this difference was not statistically significant, providing some support for the hypothesis that H. atra in the LDT exhibit a higher fission rates. At the end of the experiment H. atra in LDT were significantly longer and heavier than in HDT. H. atra surpassed their initial deployment weight and length after 13 months in the LDT by 115.2% and 45.2% respectively and in the HDT 86.9% and 24.6% respectively, changing from the small to the large phenotype known for this species. This differential growth may be linked to habitat stability and high benthic productivity and demonstrates the phenotypic plasticity of holothuroids and potential to achieve ‘Optimum Individual Size’ with respect to environmental conditions. Our results will assist in finetuning conceptual models on asexual reproduction and future experimental studies on the phenomena of fission and plastic growth in holothuroids.     

Regeneration and asexual reproduction share common molecular changes: upregulation of a neural glycoepitope during morphallaxis in Lumbriculus

Neural morphallaxis is a regenerative process characterized by wide-spread anatomical and physiological changes in an adult nervous system. During segmental regeneration of the annelid worm, Lumbriculus variegatus, neural morphallaxis involved a reorganization of sensory, interneuronal, and motor systems as posterior fragments gained a more anterior body position. A monoclonal antibody, Lan 3-2, which labels a neural glyco-domain in the leech, was reactive in Lumbriculus. In the worm, this antibody labeled neural structures, particularly axonal tracts and giant fiber pathways of the central nervous system. A 60kDa protein, possessing a lumbriculid mannose-rich glycoepitope, was upregulated during neural morphallaxis, peaking in its expression at 3 weeks post-amputation. Peak upregulation of the Lan 3-2 epitope, or the protein possessing it, corresponded to the time of major neurobehavioral plasticity during regeneration. Analyses of asexually reproducing animals also revealed induction of the Lan 3-2 epitope. In this developmental context, Lan 3-2 epitope upregulation was also confined to segments expressing both changes in positional identity and neurobehavioral plasticity, but these molecular and behavioral changes occurred prior to body fragmentation. These results suggest that the lumbriculid Lan 3-2 glycoepitope and proteins that bear them have been co-opted for neural morphallactic programs, induced both in anticipation of reproductive fragmentation and in compensation for injury-induced fragmentation.     

Latitudinal trends in reproduction,recruitment and population characteristics of some rocky littoral molluscs and cirripedes

This paper reviews progress in the COST 647 rocky littoral programme involving three patellids, two trochids and two cirripedes on European Atlantic coasts. Northern geographical limits are set primarily by repopulation failure, and northern populations are characterised by short, mid-summer breeding periods, high frequency of failed or poor recruitments and low density of large, long-lived individuals. Towards the south molluscan breeding periods lengthen and become later, extending in some cases throughout much of the year but with least or no activity in mid-summer. There are probably fewer recruitment failures and thus higher densities, but individuals are of smaller maximum size and shorter life-span.The cirripedes show similar latitudinal trends in recruitment timing but Semibalanus is restricted to a single annual brood throughout its range.The two species reaching their southern limits show progressive restriction to the lowest tidal levels.Recruitment failures do not result from inadequate gonad activity. They arise in cirripedes during the planktonic phase and in the molluscs during settlement and early shore life. The temperature sensitivity of molluscan spat is primarily responsible for the north/south gradient in recruitment times from summer towards winter.Formerly University of Leeds, England     

Seasonal variation in growth of greater snow goose goslings: the role of food supply

Even though growth rate is an important fitness component, it is still controversial to what extent parent birds adjust the timing of offspring hatch to natural variations in food supply to maximize offspring growth. We studied the role of food availability in explaining inter- and intra-seasonal variation of growth rate in goslings of greater snow geese over 5 years. The peak of hatching generally coincided with the peak of food availability. However, early-hatched goslings usually grew faster than birds hatched at the peak, which in␣turn grew faster than late-hatched goslings, although this phenomenon was not observed in all years. There was considerable variation in growth rate among the five years, the smallest goslings produced in the best year (1991) being larger than the largest goslings of the poorest year (1994). We developed three indices of food availability, based on the cumulative availability of plant biomass and nitrogen content during the growth period, and showed that the cumulative exposure to nitrogen biomass explained up to 43% of variation (intra- and inter-annual) in body size just before fledging. In years with good feeding conditions, early-hatched goslings had access to more nitrogen during their growing period than those hatching on or after the peak and they grew faster. In years of lower food availability, early-hatched goslings had no detectable advantage over peak- or late-hatched birds for access to protein-rich food and no seasonal decline in growth rate was observed. These results confirm the critical role of food supply in the seasonal variation of growth rate in Arctic-nesting geese. Received: 27 June 1997 / Accepted: 9 October 1997     

Photosynthetic plasticity in Potamogeton pectinatus L. from Argentina: strategies to survive adverse light conditions

Argentine Potamogeton pectinatus L. was grown in The Netherlands under laboratory conditions at four light intensities (50, 100, 150 and 200 µE m^–2 s^–1), and photosynthetic performance was evaluated after about 1, 2 and 3 months of growth. At these moments, chlorophyll-a and -b and tissue N and P content were also determined. During the growing period, plant lengths and number of secondary shoots were measured. In the field in Argentina, photosynthetic performance of P. pectinatus was also measured at different light intensities created by artificial shading at various times during the growing season. Field and laboratory photosynthetic results were in good agreement. P. pectinatus showed a significant plasticityin its photosynthesis, rather than in morphology. A fairly constant maximum photosynthetic rate with reduced light enabled the plants to maintain netproduction rates rather unaffected at low light intensities. Still, it can be predicted that increasing turbidity from 1–2 m^–1 at present to 3 m^–1 could lead to a strongly light-limited growth which should reduce the present weed problem considerably. Such a turbidity increase might be achieved by the introduction of a fairly dense bottom-feeding fish population like Common carp (Cyprinus carpio L.).     

放牧影响下羊草种群生殖生态学的研究

在种群水平上系统研究了放牧对松嫩草原关草种群无性繁殖、有性生殖和生物量的生殖分配的影响。结果表明,放牧制约了羊草种群无性繁殖和有性生殖的更新途径,表现在随放牧强度增加,无性繁殖体营养技密度、根茎牙密度和有性生殖体种子生物量、结实数和生殖枝分化率等指标均显著下降,是生牧后下降迅速。随放牧强度增加,关草种群根茎生物量的分配比例显著增加,而同化器官、非同化器官和种子生物量分配比例都显著下降,极牧阶段几科     

Simulated sea level change alters anatomy, physiology, growth, and reproduction of red mangrove (Rhizophora mangle L.)

Tropical coastal forests – mangroves – will be one of the first ecosystems to be affected by altered sea levels accompanying global climate change. Responses of mangrove forests to changing sea levels depend on reactions of individual plants, yet such responses have not been addressed experimentally. We report data from a long-term greenhouse study that assessed physiological and individual growth responses of the dominant neotropical mangrove, Rhizophora mangle, to levels of inundation expected to occur in the Caribbean within 50–100 years. In this study, we grew potted plants in tanks with simulated semidiurnal (twice daily) high tides that approximated current conditions (MW plants), a 16-cm increase in sea level (LW plants), and a 16-cm decrease in sea level (HW plants). The experiment lasted 2½ years, beginning with mangrove seedlings and terminating after plants began to reproduce. Environmental (air temperature, relative humidity, photosynthetically active radiation) and edaphic conditions (pH, redox, soil sulfide) approximated field conditions in Belize, the source locale for the seedlings. HW plants were shorter and narrower, and produced fewer branches and leaves, responses correlated with the development of acid-sulfide soils in their pots. LW plants initially grew more rapidly than MW plants. However, the growth of LW plants slowed dramatically once they reached the sapling stage, and by the end of the experiment, MW plants were 10–20% larger in all measured growth parameters. Plants did not exhibit differences in allometric growth as a function of inundation. Anatomical characteristics of leaves did not differ among treatments. Both foliar C:N and root porosity decreased from LW through MW to HW. Relative to LW and HW plants, MW plants had 1–7% fewer stomata/mm², 6–21% greater maximum photosynthetic rates, 3–23% greater absolute relative growth rates (RGRs), and a 30% higher RGR for a given increase in net assimilation rate. Reduced growth of R. mangle under realistic conditions approximating future inundation depths likely will temper projected increased growth of this species under concomitant increases in the atmospheric concentration of CO₂.     

Escaping an ecological dead-end: asexual overwintering and morph determination in the lettuce root aphid Pemphigus bursarius L.

1. Pemphigus bursarius exhibits a host-alternating life-cycle, overwintering as eggs on the primary host plant (poplar) before migrating in summer to the secondary host plant (mainly annual Compositae). A proportion of the population does not produce return migrants (sexuparae) in the autumn but remains in the soil and overwinters as asexual apterae, even after the annual plants have died in early winter. Laboratory and field experiments investigated the biology and overwintering success of these asexual apterae. 2. Both temperature and photoperiod are important in morph determination. The highest number of apterae (approximately 50% of progeny) was produced in a 20 °C, LD 18 : 6 h regime. Very few apterae were produced in an alatae-inducing environment (15 °C, LD 12 : 12 h), but the proportion increased over successive generations. 3. Apterae remaining in the soil in the autumn overwintered successfully in large numbers and were able to reinfest directly the root systems of newly planted lettuce grown in the same field in the following growing season. 4. Overwintered asexual populations also produced alates in July, which were able to colonise other lettuce plants, indicating that they were not sexuparae. Hence, P. bursarius can avoid the ecological dead-end that would occur through local patch extinction. Clones can therefore persist indefinitely as both asexual apterae and alatae without the need to return to poplar and undergo the sexual phase of the life-cycle.     

Latitudinal variation in thermal ecology of North American ratsnakes and its implications for the effect of climate warming on snakes

Behavioral thermoregulation is expected to be critical in determining the capacity of reptiles to respond to climate warming and how that response will vary with latitude. We used radio-telemetry to compare behavioral thermoregulation among ratsnake (Elaphe obsoleta) populations in Texas, Illinois, and Ontario, a latitudinal distance of >1500 km. Despite numerous specific differences among populations, overall the thermal ecology was surprisingly similar during the months that snakes in all three populations were active. Preferred temperatures varied only slightly across the snakes’ range, the extent of thermoregulation was similar, and by varying when during the day and season they thermoregulated, snakes in all three populations realized body temperatures within their preferred temperature range 15–20% of the time. The ability to use fine-scale behavioral thermoregulation (i.e., selective use of habitats and microclimates) to a similar extent and achieve similar outcomes across such a wide latitudinal and climatic gradient is made possible by large-scale differences in timing of activity (ratsnakes in Texas switch to nocturnal activity during summer, whereas in Illinois and Ontario activity is exclusively diurnal and hibernation lasts 5–7 months). Modeling indicated that a 3 °C increase in ambient temperature will generally improve thermal conditions for all three populations. Our empirical analyses suggest that the snakes’ ability to respond to climate warming will be determined more by their capacity to adjust when they are active than by changes in the extent of fine-scale behavioral thermoregulation. The ability to adjust timing of activity appears to make many snakes fundamentally different from lizards. As such, the consequences of climate warming may be very different for these two groups of reptiles.     

Sexual and vegetative reproduction of Hieracium pilosella L. under competition and disturbance: a grid-based simulation model

We used a spatially explicit simulation model to examine the relative importance of vegetative and sexual reproduction in Hieracium pilosella L. Based on an understanding of the complex life cycle of this species and on data from in situ population dynamics in a calcareous grassland in NW Switzerland, we simulated growth and the relative contribution of clonal reproduction by stolons and reproduction by seeds across a gradient of increasing soil fertility. Competition by a clonal grass resulted in nearly complete exclusion of H. pilosella from the more fertile part of the simulation plot. Under low soil fertility, when grass could not survive, H. pilosella largely persisted by vegetatively produced rosettes. This pattern of a sharp separation of both species was shifted slightly in favour of H. pilosella by introducing random disturbances. Only by adding: (1) long-distance seed dispersal, and (2) facilitation of seedling establishment in the vicinity of grass tussocks in vegetation gaps was a more realistic representation of field observations realised, with rosettes of H. pilosella grown from seeds occasionally distributed within dense grass vegetation. Phenotypic plasticity of stolon length was a decisive factor for the maintenance of H. pilosella populations. We conclude that a mixed strategy of clonal growth and reproduction by seeds in H. pilosella is necessary to maintain populations of this species in the presence of high interspecific competition and a shortage of open space.     

Effects of waterfowl, large fish and periphyton on the spring growth of Potamogeton pectinatus L. in Lake Mogan, Turkey

It has been argued that waterfowl and fish may threaten growth of submerged macrophytes, especially in spring during the early growth phase when plant biomass is low. A small reduction of biomass at that time might delay growth or decrease subsequent productivity. We investigated the impact of waterfowl and large fish on the spring growth of fennel pondweed (Potamogeton pectinatusL.) by employing an exclosure experiment in the macrophyte-dominated clear-water Lake Mogan, Turkey. Birds and large fish were excluded from eight plots and both in situvegetation and macrophytes kept in pots were compared to eight open plots. Also, to investigate the effect of periphyton on plant growth it was removed from half of the pot plants. Exclusion of waterfowl and fish may decrease predation on macroinvertebrates, which in turn may affect periphyton, and macrophyte growth, why macroinvertebrates also were sampled. Waterfowl density was high (15–70 ind. of coot, Fulica atraL. ha^–1), abundance of submerged plants was also high with a surface coverage of 70–80%, and benthivorous fish were present, mainly tench, (Tinca tincaL.) and carp, (Cyprinus carpioL.). Exclusion of waterfowl and large fish did not significantly affect the spring growth of pondweed; neither plants growing in situnor kept in pots. Removal of periphyton from the plants in the pots did not favour growth. The density of macroinvertebrates was not affected by the exclusion of waterfowl and large fish, but it was positively related to aboveground biomass of fennel pondweed. We suggest that even if waterfowl and large fish are in high densities, their effect on fennel pondweed spring growth in lakes with abundant submerged vegetation, such as Lake Mogan, is low.     

Recovery of Potamogeton pectinatus L. stands in a shallow eutrophic lake under extreme grazing pressure

In shallow lakes, submerged macrophytes contribute to the stabilization of the clear water state. If lost, a number of mechanisms prevent re-colonization. Lake Müggelsee (730 ha) lost its submerged vegetation due to increasing eutrophication and switched to phytoplankton dominance in 1970. After the reduction of nutrient loading in 1990, Potamogeton pectinatus L. started re-colonizing the lake. During the following years, it spread at a mean rate of 2.5 ha per year to all available areas <80 cm depth. Between 1993 and 1999, decreasing maximum biomass indicated hampered growth. Exclosure experiments revealed that herbivory reduced the aboveground biomass by more than 90%. Both waterfowl and fish were found to contribute to the grazing pressure despite a low abundance of the known herbivorous fish species and waterfowl in spring and summer. Protection of stands against grazing resulted in higher biomass of shoots, whereas shoot and tuber density did not change. Both shading by phytoplankton and periphyton, as well as grazing pressure, prevented the submerged vegetation of Lake Müggelsee from developing back to a dense zone that contributed to the reduction of turbidity.     

Latitudinal variation in intertidal algal community structure: the influence of grazing and vegetative propagation

Summary The hypothesis that sea urchin grazing and interactions with turf-forming red algae prevent large brown algae from forming an extensive canopy in the low intertidal zone of southern California was tested with field experiments at two study sites. Experimental removal of sea urchins resulted in rapid algal recruitment. Crustose coralline algae which typically dominate the substratum in areas with dense urchin populations were quickly overgrown by several species of short-lived green, brown and red algae. The removal of urchins also significantly increased the recruitment of two long-lived species of large brown algae (Egregia laevigata and Cystoseira osmundacea at one study site and E. laevigata and Halidrys dioica at the other). The experimental plots at both sites were eventually dominated by perennial red algae.A two-factorial experiment demonstrated that sea urchin grazing and preemption of space by red algae in areas where urchins are less abundant are responsible for the rarity of large brown algae in the low intertidal of southern California. The three dominant perennial red algae, Gigartina canaliculata, Laurencia pacifica and Gastroclonium coulteri, recruit seasonally from settled spores but can rapidly fill open space with vigorous vegetative growth throughout the year. These species encroach laterally into space created by the deaths of large brown algae or by other disturbances. Once extensive turfs of these red algae are established further invasion is inhibited. This interaction of algae which proliferate vegetatively with algae which recruit only from settled spores is analogous to those which occur between solitary and colonial marine invertebrates and between solitary and cloning terrestrial plants.It is suggested that a north-south gradient in the abundance of vegetatively propagating species, in grazing intensity and in the frequency of space-clearing disturbances, may account for latitudinal variation in intertidal algal community structure along the Pacific coast of North America.     

Optimal allocation to vegetative and sexual reproduction in plants: the effect of ramet density

Vegetative reproduction is a very common alternative by which plants can contribute to the next generations. There are many considerations predicting which mode of reproduction, vegetative or sexual, should be favored and numerous experimental studies to verify them. However, the results are inconsistent especially when the effect of plant density is considered. I apply here a dynamic optimization model to predict the rate of vegetative and sexual reproduction in plants as a response to changes in the local plant density. The population is assumed to occupy a heterogeneous environment consisting of patches in which growth and reproduction of plants are possible and unfavorable space between them. As the environment is globally stable, the seeds, which can disperse without restriction, exhibit a constant recruitment rate. The ramets are assumed to settle only within the patch of the mother plant. The rate of ramet production effects local density, which in turn determines ramet recruitment. The optimal strategy maximizes the expected lifetime genetic contribution, realized via both vegetative and sexual reproduction. The solutions obtained under these assumptions are dualistic. The model predicts that different approaches applied in studying the effect of ramet density should give opposite outcomes. When the comparison is between patches in natural populations, a positive relationship between relative ramet allocation and density is expected. When the density is experimentally manipulated or its effect is analyzed across different successional stages, a negative relationship should be found. The results seem to be confirmed by empirical studies.     

Water relations,gas exchange,and growth of resprouts and mature plant shoots of Arbutus unedo L. and Quercus ilex L.

Resprout and mature plant shoot growth, leaf water status and gas exchange behavior, tissue nutrient content, flowering, and production were studied for co-occurring shallow-rooted (Arbutus unedo L.) and deeprooted (Quercus ilex L.) Mediterranean tree species at the Collserola Natural Park in Northeast Spain Resprouts showed higher growth rates than mature plant shoots. During fall, no differences in eco-physiological performance of leaves were found, but mobilization of carbohydrates from burls strongly stimulated growth of fall resprouts compared to spring resprouts, despite low exposed leaf area of the fall shoots. During summer drought, resprouts exhibited improved water status and carbon fixation compared to mature plant shoots. Shoot growth of Q. ilex was apparently extended due to deep rooting so that initial slower growth during spring and early summer as compared to A. unedo was compensated. Tissue nutrient contents varied only slightly and are postulated to be of minor importance in controlling rate of shoot growth, perhaps due to the relatively fertile soil of the site. Fall flowering appeared to inhibit fall shoot growth in A. unedo, but did not occur in Q. ilex. The results demonstrate that comparative examinations utilizing vegetation elements with differing morphological and physiological adaptations can be used to analyze relatively complex phenomena related to resprouting behavior. The studies provide an important multi-dimensional background framework for further studies of resprouting in the European Mediterranean region.