Responses to simulated herbivory and water stress in two tropical C4 grasses

Summary The African grass Hyparrhenia rufa has established itself successfully in South American savannas (Llanos) and displaced dominant native grasses such as Trachypogon plumosus from the wetter and more fertile habitats. Several ecophysiological traits have been related to the higher competitive capacity of H. rufa. To further analyze the behavior of both species, their growth, biomass allocation, physiological and architectural responses to defoliation and water stress were compared under controlled conditions. Although total, aerial and underground biomass decreased under defoliation in both grasses, increases in clipped-leaf biomass and area compensated for defoliation in H. rufa but not in T. plumosus. This difference was due mainly to a higher proportion of assimilates being directed to leaf and tiller production and a higher leaf growth rate in the African grass as compared to T. plumosus, which showed incrased senescence under frequent defoliation. In both species, water stress ameliorated the effects of defoliation. The ability to compensate for defoliated biomass in H. rufa is possibly related to its long coevolution with large herbivores in its original African habitat and is apparently one of the causes of its success in Neotropical savannas.     

Evaluation of intraguild interactions between two species of insect herbivores on Pistia stratiotes

Pistia stratiotes L. (Araceae) is an important weed in many waterways around the world. The South American weevil Neohydronomus affinis Hustache (Coleoptera: Curculionidae) is a successful biocontrol agent for this weed but additional agents are needed for some areas. The planthopper Lepidelphax pistiae Remes Lenicov (Hemiptera: Delphacidae) is a specific herbivore of P. stratiotes and is highly damaging in laboratory conditions. A field experiment was designed to evaluate the damage potential of L. pistiae compared to N. affinis and to assess the potential for competition between the two herbivores. Both herbivore species were reared inside floating cages on a P. stratiotes-infested lake. Plant growth indices assessed were clonation levels, biomass and plant diameter. The growth indices of plants exposed to the herbivores, individually and combined, were comparable, and significantly lower than in the control treatments. Population levels of N. affinis inside the cages were not significantly different when alone or together with L. pistae. Conversely, the density of L. pistiae was significantly lower in combination with the weevil. This suggests that L. pistiae is a damaging herbivore that will not reduce the biological control effectiveness of N. affinis.     

Combined effects of enhanced UV-B radiation and additional nutrients on growth of two Mediterranean plant species

Seedlings of two Mediterranean plants, the slow-growing, evergreen sclerophyll Ceratonia siliqua L. and the fast growing drought semi-deciduous Phlomis fruticosa L., were grown for one year in the field at ambient or ambient plus supplemental UV-B radiation (equivalent to a 15% ozone depletion) and two levels of applied fertilizers (NPK). The effects on growth, morphological, anatomical and physiological parameters were measured at final plant harvest. Additional nutrients increased leaf nitrogen, improved growth and reduced the root/shoot ratio in both plants, yet these effects were more pronounced in the fast growing P. fruticosa. A nutrient-induced increase in chlorophyll content was also observed in this plant. The growth responses to UV-B radiation were different for the two species. Growth in C. siliqua was not affected by UV-B radiation at both nutrient levels and the same was true for P. fruticosa at low nutrients. However, at the high nutrient level, supplemental UV-B radiation improved growth in P. fruticosa, indicating a strong interaction between the treatments. Photosystem II (PSII) photochemical efficiency, methanol-extractable UV-B absorbing capacity, total phenolics and tannins were not affected by either treatment in both plants. It is concluded that nutrient levels can strongly modify the UV-B radiation effects on growth of P. fruticosa. We presume that this may be correlated to the fast growing habit of this species.     

Facilitative interactions between two lepidopteran herbivores of Asimina

Summary Insect herbivores that require young foliage for successful larval development are often restricted to a single generation during a year by the scarcity of suitable food over most of the growing season. The major specialist herbivore attacking shrubs in the genus Asimina in Florida, Eurytides marcellus, requires young foliage for successful larval development. Field manipulations were used to investigate the role of the young foliage produced by Asimina in response to defoliation by the late-season feeder Omphalocera munroei, a second specialist herbivore of Asimina in Florida, in maintaining Eurytides populations during the summer months when young foliage is otherwise scarce. Defoliation by Omphalocera proved to be the major inducer of young growth during the summer because Omphalocera defoliated Asimina shrubs so frequently and severely. When compared to young leaves produced in the absence of damage, the teaves produced by Asimina in response to defoliation were equally as suitable as food for Eurytides larvae and as acceptable as oviposition sites by Eurytides females. The availability of young foliage in an Asimina population was correlated with the size of the associated Eurytides population. The combination of regular, severe defoliation by Omphalocera and lack of a defensive response to damage by Asimina lead to a positive affect of Omphalocera on Eurytides population size, and may be central to other facilitative interactions between herbivores as well.     

Pathogen-activated induced resistance of cucumber: response of arthropod herbivores to systemically protected leaves

Summary Restricted (non-systemic) inoculation of cucurbits, green bean, tobacco, and other plants with certain viruses, bacteria, or fungi has been shown to induce persistent, systemic resistance to a wide range of diseases caused by diverse pathogens. The non-specificity of this response has fueled speculation that it may also affect plant suitability for arthropod herbivores, and there is limited evidence, mainly from work with tobacco, which suggests that this may indeed occur. Young cucumber plants were immunized by restricted infection of a lower leaf with tobacco necrosis virus (TNV), and upper leaves were later challenged with anthracnose fungus, Colletotrichum lagenarium, to confirm induction of systemic resistance to a different pathogen. The response of arthropod herbivores was simultaneously measured on non-infected, systemically protected leaves of the same plants. As has been reported before, immunization with TNV gave a high degree of protection from C. lagenarium, reducing the number of lesions and the area of fungal necrosis by 65–93%. However, there was no systemic effect on population growth of twospotted spider mites, Tetranychus urticae Koch, on upper leaves, nor did restricted TNV infection of leaf tissue on one side of the mid-vein systemically affect mite performance on the opposite, virus-free side of the leaf. Similarly, there were no effects on growth rate, pupal weight, or survival when fall armyworm larvae were reared on systemically protected leaves from induced plants. In free-choice tests, greenhouse whiteflies oviposited indiscriminately on induced and control plants. Feeding preference of fall armyworms was variable, but striped cucumber beetles consistently fed more on induced than on control plants. There was no increase in levels of cucurbitacins, however, in systemically-protected foliage of induced plants. These findings indicate that pathogen-activated induced resistance of cucumber is unlikely to provide significant protection from herbivory. The mechanisms and specificity of induced resistance in cucurbits apparently differ in response to induction by pathogens or herbivores.     

Effect of water stress on growth, Na+ and K+ accumulation and water use efficiency in relation to osmotic adjustment in two populations of Atriplex halimus L.

The effect of water stress on growth, Na⁺ and K⁺ accumulation and water utilization was investigated in plants of two populations of Atriplex halimus L. originating from Kairouan (Tunisia) and Tensift (Morocco). Water deficit was applied by withholding water for 22 days. All plants remained alive until the end of the treatment although growth was strongly reduced in both populations. Water stress decreased CO₂ assimilation in saturating conditions, mainly in the population obtained from Kairouan, suggesting an impact of drought on the dark phase of photosynthesis, beside a decrease in stomatal conductance which was recorded mainly in the population obtained from Tensift. The two studied populations did not differ in their water consumption, as indicated by similar soil gravimetric water content and plant transpiration. However, water use efficiency increased under stress conditions in the population from Tensift but not in the population from Kairouan. Thelatter population displayed a larger capacity for osmotic adjustment. A drought-induced specific increase in Na⁺ concentration was also reported in both populations. It is concluded that in A. halimus, water stress resistance estimated in terms of biomass production, could be associated with higher WUE rather than with with a greater osmotic adjustment and that sodium may assume a specific physiological function in this xerohalophytic C₄ species.     

Long-term suppression of insect herbivores increases the production and growth of Solidago altissima rhizomes

Summary Although insect herbivores have many well documented effects on plant performance, there are few studies that assess the impact of above-ground herbivory on below-ground plant growth. For a seven year period in which no large-scale herbivore outbreaks occurred, a broad spectrum insecticide was utilized to suppress herbivorous insects in a natural community dominated by Solidago altissima. Ramet heights, rhizome lengths, rhizome biomass, and the number of daughter rhizomes all were lower in the control plots than in the insecticidetreated plots. These effects should lead to a decrease in the fitness of genets in the control plots relative to the fitness of genets in the insecticide-treated plots. We also found that ramets in the control plots appear to have compensated for herbivory: the ratio of rhizome length to rhizome biomass was greatest in the control plots, which indicates that clones moved farther per unit biomass in these plots than in the insecticide-treated plots. Clonal growth models show that this shift in allocation patterns greatly reduced the magnitude of treatment differences in long-term clonal displacements.Previous work has shown, and this study verified, that clonal growth in S. altissima is well represented by random-walk and diffusion models. Therefore, we used these models to examine possible treatment differences in rates of clonal expansion. Although rhizome lengths were greater in the insecticide-treated plots, results from the models suggest that our treatments had little impact on the short- and long-term displacement of S. altissima ramets from a point of origin. This occurred because S. altissima ramets backtrack often, and thus, treatment differences in net displacements are less pronounced than treatment differences in rhizome lengths.     

Hypericum perforatum plant cells reduce Agrobacterium viability during co-cultivation

Plant recalcitrance is the major barrier in developing Agrobacterium-mediated transformation protocols for several important plant species. Despite the substantial knowledge of T-DNA transfer process, very little is known about the factors leading to the plant recalcitrance. Here, we analyzed the basis of Hypericum perforatum L. (HP) recalcitrance to Agrobacterium-mediated transformation using cell suspension culture. When challenged with Agrobacterium, HP cells swiftly produced an intense oxidative burst, a typical reaction of plant defense. Agrobacterium viability started to decline and reached 99% mortality within 12 h, while the plant cells did not suffer apoptotic process. This is the first evidence showing that the reduction of Agrobacterium viability during co-cultivation with recalcitrant plant cells can affect transformation.     

Impact of two specialist insect herbivores on reproduction of horse nettle,Solanum carolinense

The frequency of coevolution as a process of strong mutual interaction between a single plant and herbivore species has been questioned in light of more commonly observed, complex relationships between a plant and a suite of herbivore species. Despite recognition of the possibility of diffuse coevolution, relatively few studies have examined ecological responses of plants to herbivores in complex associations. We studied the impact of two specialist herbivores, the horse nettle beetle, Leptinotarsa juncta, and the eggplant flea beetle, Epitrix fuscula, on reproduction of their host, Solanum carolinense. Our study involved field and controlled-environment experimental tests of the impact on sexual and potential asexual reproduction of attack by individuals of the two herbivore species, individually and in combination. Field tests demonstrated that under normal levels of phytophagous insect attack, horse nettle plants experienced a reduction in fruit production of more than 75% compared with plants from which insects were excluded. In controlled-environment experiments using enclosure-exclosure cages, the horse nettle's two principal herbivores, the flea beetle and the horse nettle beetle, caused decreases in sexual reproduction similar to those observed in the field, and a reduction in potential asexual reproduction, represented by root biomass. Attack by each herbivore reduced the numbers of fruits produced, and root growth, when feeding in isolation. When both species were feeding together, fruit production, but not root growth, was lower than when either beetle species fed alone. Ecological interactions between horse nettle and its two primary herbivores necessary for diffuse coevolution to occur were evident from an overall analysis of the statistical interactions between the two herbivores for combined assessment of fruit and vegetative traits. For either of these traits alone, the interactions necessary to promote diffuse coevolution apparently were lacking.     

Flooding effects on starch partitioning during early growth of two oak species

Pedunculate (Quercus robur L.) and sessile (Q. petraea [Matt.] Liebl.) oaks are the most common oak species in Western Europe. They are known to display different ecological requirements, particularly relative to root hypoxia induced by flooding: In a glasshouse study of seedlings, we quantified the effects of flooding on starch mobilization from cotyledons and starch partitioning. Growth and distribution of lateral roots were also measured. The above-ground growth of Q. robur was less affected by flooding than that of Q. petraea which failed to develop a second flush. Root growth was also severely inhibited, particularly in Q. petraea. In Q. robur, lateral root initiation as well as elongation was restricted to the soil surface layer. Flooding markedly reduced total growth and concentrations of in all components except stems. Starch mobilization from cotyledons was delayed by flooding, especially in Q. robur seedlings. Under flooding, the decrease of cotyledons dry mass and starch content in Q. robur was lower than in Q. petraea, whereas Q. robur displayed larger growth than Q. petraea. The features of carbohydrate management may be crucial in the observed differences in flooding tolerance of these species.     

Combined effects of endo- and exogenous trehalose on stress tolerance and biocontrol efficacy of two antagonistic yeasts

Effects of endo- and exogenous trehalose on viability of two antagonistic yeasts, Cryptococcus laurentii (Kuffer.) Skinner and Rhodotorula glutinis (Fresen.) Harrison, were investigated after being treated with rapid-freezing, slow-freezing and freeze-drying, respectively. The accumulation of intracellular trehalose in the two yeasts was induced by culturing the yeast cells in trehalose-containing medium, which significantly enhanced viabilities of both yeasts in the slow-freezing test. Trehalose, as an exogenous protectant, at the concentration of 5% or 10% could markedly increase survivals of the two yeasts when subjected to freeze-drying. When combined with exogenous trehalose as a protective substance, the yeasts containing high intracellular trehalose level showed higher viabilities as compared to those containing low levels under both freezing and freeze-drying stresses. The highest survival of C. laurentii and R. glutinis were 90% and 97% after freeze-drying, respectively, compared to 63% and 28% for the yeasts with lower intracellular trehalose levels. These results may be due to the fact that a combined effect occurred between endo- and exogenous trehalose of yeast cells. The combined effect on C. laurentii and R. glutinis also resulted in the highest level of biocontrol efficacy against blue mold in apple fruit caused by Penicillium expansum Link, and reduced the disease indexes to 45 and 56, respectively, compared to 94 and 81 in the untreated control. Meanwhile, the combination of endo- and exogenous trehalose significantly increased population of both yeasts in apple wounds, especially at the first 48 h after inoculation, which might explain the reason of the improvement in biocontrol effects of the two yeasts.     

Use of simultaneous analysis of gas-exchange and chlorophyll fluorescence quenching for analysing the effects of water stress on photosynthesis in apple leaves

Summary A convenient system for the rapid simultaneous measurement of both chlorophyll fluorescence quenching using a modulated light system, and of CO₂, and water vapour exchange by leaves is described. The system was used in a study of the effects of water deficits on the photosynthesis by apple leaves (Malus x domestica Borkh.). Apple leaves were found to have low values of steady-state variable fluorescence, and the existence of significant fluorescence with open traps (F_o) quenching necessitated the measurement and use of a corrected F_o in the calculation of quenching components. Long-term water stress had a marked effect on both gas-exchange and chlorophyll fluorescence quenching. Non-photochemical quenching (qn) in particular was increased in water-stressed leaves, and it was particularly sensitive to incident radiation in such leaves. In contrast, rapid dehydration only affected gas exchange. Relaxation of qn quenching in the dark was slow, taking approximately 10 min for a 50% recovery, in well-watered and in draughted plants, and whether or not the plants had been exposed to high light.     

Interacting effects of grass height and herbivores on the establishment of an encroaching savanna shrub

Shrub encroachment is a widely observed problem in Southern African savannas. Although the effects of herbivory and grass height on woody species recruitment have been studied individually, little information exists about how these factors interact. In this study seeds and seedlings of the encroaching shrub Dichrostachys cinerea were planted in clipped and unclipped grass plots, with and without large herbivores present. Seed germination, seedling survival and seedling predation were monitored for 8 months. Germination started earlier in plots where herbivores were excluded. Overall, the earlier the seeds germinated, the longer the seedlings survived. Clipping positively affected the number of germinated seeds, seedling growth and survival but effects varied among herbivore exclusion treatments and sites. Invertebrates caused the majority of the seedling damage. We conclude the recruitment of D. cinerea is influenced by the interplay of grass height and herbivory. In this study, the presence of large herbivores early in the wet season, and the absence of simulated grazing later on, affected the regeneration of D. cinerea negatively. However, differences in effects among sites suggest that the mechanisms found here may work differently in other habitats.      

Translocation and metabolism of glycine betaine by barley plants in relation to water stress

The glycine betaine which accumulated in shoots of young barley plants (Hordeum vulgare L.) during an episode of water stress did not undergo net destruction upon relief of stress, but its distribution among plant organs changed. During stress, betaine accumulated primarily in mature leaves, whereas it was found mainly in young leaves after rewatering. Well-watered, stressed, and stressed-rewatered plants were supplied with [methyl-¹⁴C]betaine (8.5 nmol) via an abraded spot on the second leaf blade, and incubated for 3 d. In all three treatments the added ¹⁴C migrated more or less extensively from the second leaf blade, but was recovered quantitatively from various plant organs in the form of betaine; no labeled degradation products were found in any organ. When 0.5 mol of [methyl-¹⁴C]betaine was applied via an abraded spot to the second leaf blades of well-watered, mildly-stressed, and stressed-rewatered plants, ¹⁴C was translocated out of the blades at velocities of about 0.2–0.3 cm/min which were similar to velocities found for applied [¹⁴C]sucrose. Heat-girdling of the sheath prevented export of [¹⁴C]betaine from the blade. When 0.5 mol [³H]sucrose and 0.5 mol [¹⁴C]betaine were suppled simultaneously to second leaf blades, the ³H/¹⁴C ratio in the sheath tissue was the same as that of the supplied mixture. After supplying tracer [¹⁴C]betaine aldehyde (the immediate precursor of betaine) to the second leaf blade, the ¹⁴C which was translocated into the sheath was in the form of betaine. Thus, betaine synthesized by mature leaves during stress behaves as an inert end product and upon rewatering is translocated to the expanding leaves, most probably via the phloem. Accordingly, it is suggested that the level of betaine in a barley plant might serve as a useful cumulative index of the water stress experienced during growth.     

Effects of two types of herbivores on the population dynamics of a perennial herb

The joint effects of multiple herbivores on their shared host plant have received increasing interest recently. The influence of herbivores on population dynamics of their host plants, especially the relative roles of different types of damage, is, however, still poorly understood. Here, we present a modelling approach, including both deterministic and stochastic matrix modelling, to be used in estimating fitness effects of multiple herbivores on perennial plants. We examined the effects and relative roles of two specialist herbivores, a pre-dispersal seed predator, Euphranta connexa, and a leaf-feeding moth, Abrostola asclepiadis, on the population dynamics and long-term fitness of their shared host plant, a long-lived perennial herb Vincetoxicum hirundinaria (Asclepiadaceae). We collected demographic data during 3 years and combined these data with the effects of natural levels of herbivory measured from the same individuals. We found that both seed predation and leaf herbivory reduced population growth of V. hirundinaria, but only very high damage levels changed the growth trend of the vigorously growing study populations from positive to negative. Demographic modelling indicated that seed predation had a greater impact on plant population growth than leaf herbivory. The effect of leaf herbivory was weaker and diminished with increasing level of seed predation. Evaluation of individual fitness components, however, suggested that leaf herbivory contributed more strongly to host plant fitness than seed predation. Our results emphasize that understanding the effects of a particular herbivore on plant population dynamics requires also knowledge on other herbivores present in the system, because the effect of a particular type of herbivory on plant population dynamics is likely to vary according to the intensity of other types of herbivory. Furthermore, evaluating herbivore impact from using individual fitness components does not necessarily reflect the long-term effects on total plant fitness.     

Effects of water stress on gas exchange and water relations of a succulent epiphyte,Kalanchoë uniflora

Summary Measurements of gas exchange, xylem tension and nocturnal malate synthesis were conducted with well-watered and droughted plants of Kalanchoë uniflora. Corresponding results were obtained with plants grown in 9 h and 12 h photoperiods. In well-watered plants, 50 to 90% of total CO₂-uptake occurred during the light period. Nocturnal CO₂-uptake and malate synthesis were higher and respiration rate was lower in old leaves (leaf pairs 6 to 10) compared to young leaves (leaf pairs 1 to 5). Within four days of drought distinct physiological changes occurred. Gas exchange during the light period decreased and CO₂-uptake during the dark period increased. Nocturnal malate synthesis significantly increased in young leaves.Respiration rate decreased during periods of drought, this decrease being more pronounced in young leaves compared to old leaves. Restriction of gas exchange during the light period resulted in a decrease of transpiration ratio from more than 100 to about 20. The difference between osmotic pressure and xylem tension decreased in young leaves, indicating a reduction in bulk leaf turgor-pressure.We conclude that both the CAM-enhancement in young leaves and the decrease of respiration rate are responsible for the increase of nocturnal CO₂-uptake during water stress. During short drought periods, which frequently occur in humid habitats, the observed physiological changes result in a marked reduction of water loss while net CO₂-uptake is maintained. This might be relevant for plant growth in the natural habitat.Abbreviations LP light period - DP dark period - CAM crassulacean acid metabolism     

Influence of induced water stress in ponderosa pine on pine sawflies

Summary Three levels of water stress were induced on pole-size ponderosa pine (Pinus ponderosa) to determine the influence of plant moisture stress on oviposition, survival, and growth of two species of pine sawfly (Neodiprion fulviceps and N. autumnalis). It was found that water stress affected oviposition and subsequent egg survival but not larval development or survival. Stress had a negative effect on early season oviposition (N. fulviceps) and a positive effect on late season oviposition (N. autumnalis). Egg hatch was different between species and years and among treatment levels. Larval development, feeding, and survival were not affected by water stress. Overall, the effect of stress was not sufficient to explain population outbreaks of sawflies. Several hypotheses are presented as possible explanations for the observed experimental results.     

Biomass allocation, relative competitive ability and water use efficiency of two dominant species in semiarid Loess Plateau under water stress

A better understanding of the growth and interspecific competition of native dominant species under water stress should aid in prediction of succession in plant communities. In addition, such research would guide the selection of appropriate conservation and agricultural utilization of plants in semiarid environments that have not been very well characterized. Biomass production and allocation, relative competitive ability and water use efficiency of one C₄ herbaceous grass (Bothriochloa ischaemum) and one C₃ leguminous subshrub (Lespedeza davurica), both important species from the semiarid Loess Plateau of China, were investigated in a pot-cultivation experiment. The experiment was conducted using a replacement series design in which B. ischaemum and L. davurica were grown with twelve plants per pot, in seven combinations of the two species (12:0, 10:2, 8:4, 6:6, 4:8, 2:10, and 0:12). Three levels of water treatments included sufficient water supply (HW), moderate water stress (MW) and severe water stress (LW). These treatments were applied after seedling establishment and remained until the end of the experiment. Biomass production and its partitioning, and transpiration water use efficiency (TWUE) were determined at the end of the experiment. Interspecific competitive indices (competitive ratio (CR), aggressiveness (A) and relative yield total (RYT)) were calculated from the dry weight for shoots, roots and total biomass. Water stress decreased biomass production of both species in monoculture and mixture. The growth of L. davurica was restrained in their mixtures for each water treatment. L. davurica had significantly (P < 0.05) greater root:shoot allocation than B. ischaemum for each water treatment and proportion within the replacement series. Aggressiveness (A) values for B. ischaemum with respect to L. davurica were negative only at the proportions of B. ischaemum to L. davurica being 8:4 and 10:2 in LW treatment. B. ischaemum had a significantly (P < 0.05) higher CR value under each water treatment, and water stress considerably reduced its relative CR while increased that of L. davurica. RYT values of the two species indicated some degree of resource complimentarity under both water sufficient and deficit conditions. The results suggest that it is advantageous for growing the two species together to maximize biomass production, and the suggested ratio was 10:2 of B. ischaemum to L. davurica because of significantly higher (P < 0.05) RYT and TWUE under low water availability condition.     

Density- and growth stage-dependent responses to defoliation in two rhizomatous grasses

Summary Responses to defoliation were studied in two tallgrass prairie perennials (Andropogon gerardii and Panicum virgatum) established from seed at three densities. P. virgatum was also grown from transplanted rhizomes of established clones. Plants of both species displayed a continuum of responses to defoliation, from large reductions in biomass, tillering and seed production to significant increases in one or more performance measures. In crowded populations, defoliation shifted plants into subordinate positions within the competitive hierarchy. Plants competing intraspecifically and those that were initially small suffered more from defoliation than either plants grown at low density or those that were larger than their neighbors. At the highest plant density, the effects of defoliation or initial plant size were overshadowed by the effects of crowding. When defoliated and grown at similar densities, P. virgatum and A. gerardii grown from seed showed large reductions in biomass, seed production, and new rhizome production, but established P. virgatum ramets grown from rhizomes showed increases in these performance measures. Thus, herbivory may be particularly detrimental to P. virgatum during juvenile stages before perennating organs have developed. Overcompensation of P. virgatum clones in response to defoliation only occurred if all ramets within the clone were defoliated. In clones containing both defoliated and undamaged ramets, there were no differences in their performance, suggesting that genets are capable of integrating the effects of differential defoliation among shoots. Defoliated P. virgatum clones allocated a smaller fraction of their total biomass to new rhizomes, indicating that the short-term regrowth response following defoliation may incur a longer-term cost associated with gradual reduction in biomass of the perennating organs and reduced genet success.     

Effect of water stress on growth, osmotic adjustment, cell wall elasticity and water-use efficiency in Spartina alterniflora

In the northern spring–summer season of 2004–2005, vegetative propagated plants of Spartina alterniflora were grown under control and water stress conditions on the Mediterranean sea shore of the south-east of Tunis. Control plants were irrigated every week and water stress plants were irrigated until the soil achieved 50% (mild stress) and 25% (severe stress) field capacity (FC). Dry and fresh weight at the whole plant level (g plant⁻¹), shoot to root ratio on dry and fresh weight, photosynthesis (A), transpiration rate (E), instantaneous water-use efficiency (WUE_i), leaf water potential (Ψw), leaf water content (WC), osmotic potential at full turgor (Ψs¹⁰⁰), osmotic potential at turgor loss point (Ψs⁰), osmotic adjustment (OA), proline, sugars, inorganic compounds and cell wall elasticity (CWE) were evaluated during a period of 6 days period between 82 and 90 days after the beginning of treatment (DAT). Plants grown under severe and mild-water stress showed lower Ψw than in control plants with values that averaged −3.1, −1.6 and −0.9 MPa, respectively. S. alterniflora plants submitted to mild-water stress exhibited OA and a decrease in CWE. However, under severe water stress the OA was not observed and CWE also decreased, but it was higher than in the mild-water stress. OA was mainly explained by the accumulation of nitrates, sugars and at a lesser degree, proline. S. alterniflora had a strong decline of the dry and fresh weight of the whole plant associated to a marked decrease of photosynthesis (A) and transpiration (E) in response to water stress, although WUE_i was increased. These results suggest that OA and WUE_i can be important components of the water stress adaptation mechanism in this species, but they are not sufficient enough to contribute to resistance to water stress.