Plant biomass partitioning and chemical defense: Response to defoliation and nitrate limitation

Summary Plant growth and allocation to root, shoot and carbon-based leaf chemical defense were measured in response to defoliation and nitrate limitation inHeterotheca subaxillaris. Field and greenhouse experiments demonstrated that, following defoliation, increased allocation to the shoot results in an equal root/shoot ratio between moderately defoliated (9% shoot mass removed) and non-defoliated plants. High defoliation (28% shoot mass or >25% leaf area removed) resulted in greater proportional shoot growth, reducing the root/shot ratio relative to moderate or non-defoliated plants. However, this latter effect was dependent on nutritional status. Despite the change in distribution of biomass, defoliation and nitrate limitation slowed the growth and development ofH. subaxillaris. Chronic defoliation decreased the growth of nitrate-rich plants more than that of nitrate-limited plants. The concentration of leaf mono- and sesqui-terpenes increased with nitrate-limitation and increasing defoliation. Nutrient stress resulting from reduced allocation to root growth with defoliation may explain the greater allocation to carbon-based leaf defenses, as well as the defoliation-related greater growth reduction of nitrate-rich plants.     

Effects of manipulation of water and nitrogen regime on the water relations of the desert shrub Larrea tridentata

Summary Water and nitrogen regimes of Larrea tridentata shrubs growing in the field were manipulated during an annual cycle. Patterns of leaf water status, leaf water relations characteristics, and stomatal behavior were followed concurrently. Large variations in leaf water status in both irrigated and nonirrigated individuals were observed. Predawn and midday leaf water potentials of nonirrigated shrubs were lowest except when measurements had been preceded by significant rainfall. Despite the large seasonal variation in leaf water status, reasonably constant, high levels of turgor were maintained. Pressure-volume curve analysis suggested that changes in the bulk leaf osmotic potential at full turgor were small and that nearly all of the turgor adjustment was due to tissue elastic adjustment. The increase in tissue elasticity with increasing water deficit manifested itself as a decrease in the relative water content at zero turgor and as a decrease in the tissue bulk elastic modulus. Because of large hydration-induced displacement in the osmotic potential and relative water content at zero turgor, it was necessary to use shoots in their natural state of hydration for pressure-volume curve determinations. Large diurnal and seasonal differences in maximum stomatal conductance were observed, but could not easily be attributed to variations in leaf water potential or leaf water relations characteristics such as the turgor loss point. The single factor which seemed to account for most of the diurnal and seasonal differences in maximum stomatal conductance between individual shrubs was an index of soil/root/ shoot hydraulic resistance. Daily maximum stomatal conductance was found to decrease with increasing soil/root/ shoot hydraulic resistance. This pattern was most consistent if the hydraulic resistance calculation was based on an estimate of total canopy transpiration rather than the more commonly used transpiration per unit leaf area. The reasons for this are discussed. It is suggested that while stomatal aperture necessarily represents a major physical resistance controlling transpiration, plant hydraulic resistance may represent the functional resistance through its effects on stomatal aperture.     

Repeat flowering in apple caused by water stress or defoliation

Summary A series of experiments involving defoliation or water stress at different dates indicated that either of these treatments can make potted apple trees flower a second time in any one year, as long as the treatment is given near the end of July. The results suggest that the reflowering after a period of water stress was primarily a result of the loss of leaves that occurred when the plants were subsequently rewatered. Reflowering normally occurred only if flower primordia had already differentiated at the time of the treatment. There was an indication that in early July water stress was more effective than defoliation at stimulating reflowering.     

Controls of nitrogen limitation in tallgrass prairie

Summary The relationship between fire frequency and N limitation to foliage production in tallgrass prairie was studied with a series of fire and N addition experiments. Results indicated that fire history affected the magnitude of the vegetation response to fire and to N additions. Sites not burned for over 15 years averaged only a 9% increase in foliage biomass in response to N enrichment. In contrast, foliage production increased an average of 68% in response to N additions on annually burned sites, while infrequently burned sites, burned in the year of the study, averaged a 45% increase. These findings are consistent with reports indicating that reduced plant growth on unburned prairie is due to shading and lower soil temperatures, while foliage production on frequently burned areas is constrained by N availability. Infrequent burning of unfertilized prairie therefore results in a maximum production response in the year of burning relative to either annually burned or long-term unburned sites.Foliage biomass of tallgrass prairie is dominated by C₄ grasses; however, forb species exhibited stronger production responses to nitrogen additions than did the grasses. After four years of annual N additions, forb biomass exceeded that of grass biomass on unburned plots, and grasses exhibited a negative response to fertilizer, probably due to competition from the forbs. The dominant C₄ grasses may out-compete forbs under frequent fire conditions not only because they are better adapted to direct effects of burning, but because they can grow better under low available N regimes created by frequent fire.     

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.     

Recovery of subalpine dwarf shrub heath after neighbour removal and fertilization

We tested if subalpine heath vegetation in northern Italy recovered after experimental perturbation of soil nutrient availability (fertilization) and species composition (removal of co-dominant dwarf shrubs). Species cover was assessed non-destructively before the start of the treatments (1995), at the end of the treatments (1999) and 4 years after the treatments ended (2003). Shrub removal had rather modest effects on heath vegetation, except for mosses which decreased significantly in removal plots. Fertilization decreased the cover of shrubs, mosses, and some graminoids but increased the cover of Festuca rubra. Fertilization converted heath to grassland, but the response of graminoid species was individualistic. Fertilization decreased vascular species richness and evenness, probably through negative effects of shading and litter accumulation on plant growth or recruitment. The vegetation had not recovered completely 4 years after the perturbations had stopped. This suggests that, in contrast to rapid responses to species removal and fertilization, recovery from these perturbations was rather slow, presumably because recovery was affected by long-term biotic interactions and species controls on ecosystem properties.     

Comparative responses of the Savanna grasses Cenchrus ciliaris and Themeda triandra to defoliation

Summary Two perennial tussock grasses of savannas were compared in a glasshouse study to determine why they differed in their ability to withstand frequent, heavy grazing; Cenchrus ciliaris is tolerant and Themeda triandra is intolerant of heavy grazing. Frequent defoliation at weekly intervals for six weeks reduced shoot biomass production over a subsequent 42 day regrowth period compared with previously undefoliated plants (infrequent) in T. triandra, but not in C. ciliaris. Leaf area of T. triandra expanded rapidly following defoliation but high initial relative growth rates of shoots were not sustained after 14 days of regrowth because of reducing light utilising efficiency of leaves. Frequently defoliated plants were slower in rate of leaf area expansion and this was associated with reduced photosynthetic capacity of newly formed leaves, lower allocation of photosynthate to leaves but not lower tiller numbers. T. triandra appears well adapted to a regime where defoliation is sufficiently infrequent to allow carbon to be fixed to replace that used in initial leaf area expansion. In contrast, C. ciliaris is better adapted to frequent defoliation than is T. triandra, because horizontally orientated nodal tillers are produced below the defoliation level. This morphological adaptation resulted in a 10-fold higher leaf area remaining after defoliation compared with similarly defoliated T. triandra, which together with the maintenance of moderate levels of light utilising efficiency, contributed to the higher leaf area and shoot weight throughout the regrowth period.     

In vitro propagation of caper (Capparis spinosa L.)

Summary A twenty fold multiplication per twenty days of caper was achieved by culturing nodal shoot segments in the presence of BAP (4 μM) plus IAA (0.3 μM) and GA₃ (0.3 μM). The use of a modified MS medium facilitated this response. Plantlet regeneration was induced on single shoots taken from proliferating clusters subcultured for 20 days on a reduced BAP (2 μM) without auxin and gibberellin Higher rooting responses (70%) were obtained after a 20-day incubation period in darkness on solid half-strength MS1 medium plus IAA (30 μM), followed by a subsequent 20 day culture period on half-strength MSI basal medium. Proliferation was mainly due to axillary shoot-bud development as revealed by histological studies. The extensive meristematic activities observed indicated the enormous morphological potential of this species.     

Differences in response to defoliation between males and females of Silene dioica

Summary The response by male and female plants to herbivory was studied by experimental defoliation of the dioecious perennial herb Silene dioica in a green-house. Male and female plants were defoliated prior to and during the early flowering phase at two intensities (50% and 100% of leaf-area removed) in two consecutive years. Defoliation resulted in a decrease in the number of flowers initiated in both sexes, while a larger delay of peak flowering and a higher mortality was observed in males compared to females. In female plants, severe defoliation resulted in a reduction in seed number per capsule and in seed size compared to control. Females showed a negative correlation between the production of flowers in the first and second season in all treatments, while flowering in males the first season was not correlated with flowering in the second season. Females also showed a lower frequency of flowering than males during the two seasons studied. However, during the flowering period, males allocated significantly more biomass to flowers than did females. This outcome supports the idea that females may have a higher total reproductive expenditure than males, but males have a higher reproductive effort during flowering. Male rosette leaves were significantly preferred by the generalist herbivore Arianta arbustorum in experiments. This preference was most pronounced in trials with leaves from fertilized plants compared to nonfertilized plants. A greater storage of resources in aboveground leaves during winter by males compared to females may explain the higher preference for male leaves and the higher male mortality following early defoliation. Furthermore, males are smaller than females and may have a lower ability than females to replace lost resources needed for reproduction when defoliated early in the season.     

Intraspecific variation in the response of Themeda triandra to defoliation: the effect of time of recovery and growth rates on compensatory growth

Summary The response to a single defoliation was studied on three clones of Themeda triandra collected in the short, mid, and tall grassland regions of the Serengeti National Park (Tanzania). These sites represent a gradient of decreasing grazing intensity. Growth, allocation pattern, and several morphometric traits were monitored during an 80-day period. Clipped plants of the short and medium clones fully compensated for the reduction of biomass, while plants of the tall clone showed overcompensation. During the first two weeks after clipping, clipped plants showed lower relative growth rates than unclipped ones, whereas the opposite was observed later on. Clipped plants compensated for the removal of leaf area by producing new leaves with lower specific weights and higher nitrogen content. They also produced more, smaller tillers. Although clipped plants mobilized nonstructural carbohydrates from roots and crowns, this did not account for a significant amount of growth. Relative growth rates of unclipped plants of the short clone were higher. The relative growth rate of the short clone diminished less after clipping, but also exhibited the lowest increase later. The tall clone was the most negatively affected early, but showed the highest compensation later. Compared to the other clones, the short ecotype showed many of the characteristics that defoliation induced in each individual of any clone: higher allocation to leaf area production, higher relative growth rate, higher number but smaller size of tillers, and lower leaf specific weights.     

Flooding effects on rapid responses of the invasive plant Alternanthera philoxeroides to defoliation

In experiments under controlled growth conditions it was examined how flooding affected the responses of the invasive plant Alternanthera philoxeroides to defoliation. In drained and flooded conditions, plants were subjected to five defoliation levels: 0, 10, 50, 90% removal of leaf tissue and apex removal (90% leaf tissue plus apical bud removal). Plants were harvested weekly for five weeks. In drained conditions, plant biomasses including total biomass, shoot biomass and root biomass after 50% defoliation rapidly recovered to the control plant level. They were significantly lower for the 90% defoliation and apex removal treatments compared to control plants throughout the experiment. In flooded conditions, total biomass and shoot biomass after 50% defoliation, 90% defoliation, and apex removal treatments could return to control plant levels before the end of the experiment. In 90% defoliation and apex removal treatments root to shoot biomass ratios of both drained and flooded plants were initially much higher than in control plants, but the difference disappeared rapidly. The final biomasses decreased with increased defoliation intensity in drained conditions, but no significant difference was generally found in any of the defoliation treatments in flooded conditions. The rapid re-growth of A. philoxeroides plants after defoliation may partly be responsible for its invasion success. However, defoliation capable of removing 90% of the leaf tissue may be desirable in restricting the growth of this invasive species in drained conditions.     

The effects of nitrogen limitation on the ultrastructure of the cyanobacterium Agmenellum quadruplicatum

The effects of nitrogen limitation on the ultrastructure of the unicellular cyanobacterium, Agmenellum quadruplicatum, were studied by thin sectioning transmission electron microscopy. Nitrogen became limiting for growth 14–15 h after transfer to nitrogen-limiting medium, but cultures retained full viability for at least 45 h. The c-phycocyanin: chlorophyll a ratio and cellular nitrogen content of the culture dropped rapidly after 14–15 h, as a progressive deterioration of major cell structures took place. Phycobilisomes were degraded first, followed by ribosomes and, then, thylakoid membranes. These structures were virtually depleted from the cells within 26 h. Intracellular polysaccharide accumulated in place of the normal cell structures throughout this period. Nitrogen limitation did not affect polyphosphate bodies, carboxysomes, lipid granules, the cell envelope, or the extra-cellular glycocalyx. All of the ultrastructural changes resulting from nitrogen limitation were reversed upon addition of nitrate to a starved culture. Most cell structures were restored within 3 h, and restoration was complete within 9 h.     

Influence of temporal heterogeneity in nitrogen supply on competitive interactions in a desert shrub community

Soil nutrients in arid systems are supplied to plants in brief pulses following precipitation inputs. While these resource dynamics have been well documented, little is known about how this temporal heterogeneity influences competitive interactions. We examined the impacts of the temporal pattern of N supply on competitive intensity and ability in an N-limited desert shrub community. At our field site, the three codominant shrubs, Atriplex confertifolia, A. parryi, and Sarcobatus vermiculatus, differ in seasonal growth patterns, with A. confertifolia and S. vermiculatus achieving higher growth rates earlier in the growing season than A. parryi. We predicted that these timing differences in maximum growth rate may interact with temporal variation in N supply to alter competitive abilities over time. Seedlings of the two Atriplex species were planted either individually in field plots or as target plants surrounded by neighbor seedlings. After one year of establishment, the same amount of ¹⁵N was applied to plots either as early spring pulses, mid spring pulses or continuously through the second growing season. Competitive effects were observed under continuous and pulsed N supply. Averaged across all target–neighbor treatments, competitive intensity was ∼1.8-fold greater when N was pulsed compared to when N was supplied continuously, but overall, the outcome of competitive interactions was not influenced by N pulse timing. While the timing of resource supply did not differentially influence the competitive abilities of coexisting species in this system, the temporal pattern of resource supply did alter the intensity of competitive interactions among species. While additional studies in other systems are needed to evaluate the generality of these results, this study suggests that competitive intensity may not necessarily be a direct function of productivity or resource availability as traditionally assumed. Instead, the intensity of competitive interactions in resource-poor systems may depend upon the temporal pattern of resource supply.     

The effects of host plant defoliation and fertilizer application on larval growth and oviposition behaviour in cinnabar moth

Summary Defoliated ragwort plants produced regrowth foliage that was higher in alkaloid, but lower in amino acid concentrations than primary foliage. Total N was not affected. 2) Plants fertilized with nitrogen (as ammonium sulphate) had lower amino acid concentrations than unfertilized control plants, slightly increased alkaloid levels but similar total N concentrations. 3) Ovipositing females laid eggs upon plants with equal probability for controls, regrowth and fertilized foliage (one rosette in 5 received an egg batch). However, the probability of receiving eggs was significantly lower on the primary leaves of cut-back plants that had had their lower leaves removed a few days before egg laying (only one rosette in 13 was selected). 4) Egg batch size was higher on fertilized control foliage than on other treatments. 5) Larvae attained greater final weights when fed a diet of regrowth foliage, despite the higher levels of alkaloid they contained. Larval development rate was not affected by experimental treatment of the foliage. 6) Larval growth was lowest on the leaves of fertilized plants. This was associated with significant reductions in the concentrations of three amino acids (methionine down 29%, tyrosine 33% and lysine 25%).     

The effects of the interaction between salinity and nitrogen limitation in Agrostis stolonifera L.

Salt-tolerant Agrostis stolonifera ecotypes commonly grow on upper salt marshes, environments regarded as having a limited nitrogen supply. The interaction between salinity and nitrate supply limitation was studied in two ecotypes of A. stolonifera, one isolated from an upper salt marsh and one from an inland habitat. The ion, amino acid, glycine betaine and sugar contents of the two ecotypes were determined over a range of external salt concentrations and levels of nitrate supply. In vivo nitrate reductase activity was also measured. Several low molecular weight nitrogenous compounds accumulated in the salt-stressed plants. Nitrogen supply limitation had a great effect on the way in which the plants responded to salt stress. In particular, the concentrations of the soluble organic nitrogenous compounds were reduced. The results are discussed with respect to the salt marsh environment, and possible models for cytoplasmic osmoregulation are presented.Acknowledgements: One of us (MJH) gratefully acknowledges the receipt of a research studentship from the Science Research Council, U.K. We would also like to thank Mrs E. E. Griffiths for skilled technical assistance, and Dr I. Ahmad for help with the amino acid analysis.     

Consequences of nitrogen and phosphorus limitation for the performance of two planthoppers with divergent life-history strategies

Phytophagous insects have a much higher nitrogen and phosphorus content than their host plants, an elemental mismatch that places inherent constraints on meeting nutritional requirements. Although nitrogen limitation is well documented in insect herbivores, phosphorus limitation is poorly studied. Using factorial experiments in the laboratory and field, in which levels of soil nitrogen and phosphorus were manipulated, we studied the relative consequences of macronutrient limitation for two herbivores, namely the phloem-feeding planthoppers Prokelisia dolus and P. marginata. These planthoppers inhabit the salt marshes of North America where large stands of their Spartina host plant are found. Notably, these congeners differ in their dispersal abilities; P. marginata is dispersive whereas P. dolus is sedentary. Both nitrogen and phosphorus subsidies enhanced the nitrogen and phosphorus content of Spartina. When P. dolus and P. marginata were raised on plants with an enriched nitrogen signature, they exhibited greater survival, grew to a larger size, developed more rapidly, and achieved higher densities than on nitrogen-deficient plants. However, P. marginata experienced greater fitness penalties than P. dolus on nitrogen-deficient plants. Phosphorus limitation and associated fitness penalties were not as severe as nitrogen limitation for P. marginata, and were not detected in P. dolus. The tempered response of P. dolus to N- and P-deficient Spartina is probably due to its greater investment in feeding musculature and hence ability to compensate for nutrient deficiencies with increased ingestion. To cope with deteriorating plant quality, P. dolus employs compensatory feeding, whereas P. marginata disperses to higher quality Spartina. When its option of dispersal is eliminated and P. marginata is confined on nutrient-deficient plants, its performance is drastically reduced compared with P. dolus. This research highlights the importance of interfacing herbivore life-history strategies with ecological stoichiometry in order to interpret the consequences of macronutrient limitation on herbivore performance and population dynamics.     

Variability and ecology of the alpine dwarf shrub community Loiseleurio-Cetrarietum

Summary Variation in the dwarf shrub community Loiseleurio-Cetrarietum from Mt. Patscherkofel, Austria, was investigated using polar ordination, principal component analysis, hierarchical cluster analysis, and multiple correlation analysis. Comparison was made with existing data (relevé tables) from the Alps. In addition to phytosociological methods the canopy structure and the morphological variability of the dominant species Loiseleuria procumbens were studied. The result of the namerical phytosociological analyses yield 3 clusters representing: 1) A dense Loiseleuria carpet, 2) a Loiseleuria carpet damaged by erosion and 3) an open Loiseleuria community on skeletal soils. The first two communities can be combined as one type because of their canopy structure in the form of dense intertwined individual plants and because of the morphological uniformity of Loiseleuria, although in the second group there are species growing on erosion gaps and showing close floristic connections to the third group. The comparison of 140 relevés from the Alps proves that the Loiseleurio-Cetrarietum is characterized only by the dominance of Loiseleuria procumbens while the abundantly occurring lichens change in their species composition along the gradient of wind, and elevation. The open Loiseleuria community at the most extreme habitats (cluster 3) is regarded as a new association. Not only floristic but also physiognomical, morphological and ecophysiological evidence led to this conclusion. The name Gymnomitrio-Loiseleurietum (ass. prov.) is proposed. This plant community shows floristic and ecological correlations to the Diapensia-Loiseleuria-Cesiolichen community known from Scandinavia.Nomenclature of species follows Ehrendorfer (1973) (vascular plants), Poelt (1974) (lichens) and Gams (1973) (mosses).For PCA, hierarchical cluster analysis, and multiple correlation analysis, computer programs from the data bank system (AFO-data) of the Alpine Forschungsstelle Obergurgl, Austria were used. I am indebted to Dr. S. Pamer and P. Moser for their invaluable help. For polar ordination a separate program was developed (language: Simula, author: Dr. G. Knecht). I am also indebted to Dr. H. Hilscher for the translation of the text.     

Lipid and hydrocarbon production byBotryococcus spp. under nitrogen limitation and anaerobiosis

The production of lipids and hydrocarbons in batch cultures of the algaeBotryococcus braunii andB. protuberans has been studied with respect to nitrogen limitation under aerobic and anaerobic conditions. Nitrogen deficiency significantly decreased the dry weight, chlorophylla and protein contents but the amounts of carotenoids, carbohydrates and lipids increased in both the species. Nitrogen starvation gave a 1.6-fold increase in lipid content. Anaerobiosis under nitrogen deficient conditions gave greater lipid production than anaerobiosis in nitrogen supplemented medium. Under nitrogen deficiency, the hydrocarbon fraction increased and the polar lipids decreased. Anaerobiosis induced hydrocarbon synthesis more significantly than nitrogen deficiency but decreased other non-polar and polar lipids.