Variation in shoot mortality within crowns of severely defoliated <Emphasis Type="Italic">Betula maximowicziana</Emphasis> trees in Hokkaido,northern Japan

To clarify mortality patterns of current-year shoots within the crown of Betula maximowicziana Regel after severe insect herbivory in central Hokkaido, northern Japan, we investigated the degree of defoliation, pattern of shoot development, shoot mortality, and leaf tissue-water relations. One hundred current-year long shoots growing in a B. maximowicziana plantation were observed for defoliation and mortality in June 2002. An outbreak of herbivorous insects (Caligula japonica and Lymantria dispar praeterea) occurred in the stand in mid-to-late June, and the monitored shoots were defoliated to various degrees. Within 1 month of defoliation, some of the severely defoliated shoots had produced new leaves on short shoots that had emerged from axillary buds. Stepwise logistic regression revealed that the probability that current-year long shoots would put out axillary short shoots with leaves is closely related to the degree of defoliation. To evaluate the water relations of the leaves, we determined pressure–volume curves for the leaves that survived the herbivorous insect outbreak and the new leaves that emerged after defoliation. The water potential at turgor loss (Ψ_l,tlp) and the osmotic potential at full turgidity (Ψ_π,sat) were higher for the new leaves than for the surviving leaves, indicating a lower ability to maintain leaf cell turgor against leaf dehydration in the new leaves. Of the 100 shoots, 13 died after the emergence of new leaves. Stepwise logistic regression revealed that the probability that the long shoots would die generally increased with the emergence of new leaves, with increasing shoot height. This result suggests that the combined effect of the vulnerability of newly emerged leaves and low water availability, associated with higher shoot positions within the crown, caused shoot mortality. Based on our results, some possible mechanisms for mortality in severely defoliated B. maximowicziana are discussed.     

Defoliation increases carbon limitation in ectomycorrhizal symbiosis of <Emphasis Type="Italic">Betula pubescens</Emphasis>

Boreal forest trees are highly dependent on root-colonizing mycorrhizal fungi. Since the maintenance of mycorrhizal symbiosis implies a significant carbon cost for the host plant, the loss of photosynthetic leaf area due to herbivory is expected to reduce the host investment in mycorrhizae. We tested this hypothesis in a common garden experiment by exposing ectomycorrhizal white birch (Betula pubescens Ehrh.) seedlings to simulated insect defoliation of 50 or 100% intensity during either the previous or the current summer or repeatedly during both seasons before harvest. The shoot and root growth of the seedlings were distinctly reduced by both 100% defoliation and repeated 50% defoliation, and they were more strongly affected by previous-year than current-year defoliation. The root to shoot ratio significantly decreased after 100% defoliation, indicating reduced proportional allocation to the roots. Ergosterol concentration (i.e. fungal biomass) in the fine roots decreased by 100% defoliation conducted either in the year of harvest or in both years. No such decrease occurred following the 100% defoliation conducted in the previous year, indicating the importance of current photosynthates for fungal symbionts. The trend was similar in the colonization percentage of thick-mantled mycorrhizae in the roots, the most marked decline occurring in the repeatedly defoliated seedlings. The present results thus support the prediction that the plant investment in ectomycorrhizae may decline as a response to foliage loss. Moreover, the colonization percentage of thick-mantled mycorrhizae correlated positively with the ratio of leaf to heterotrophic plant biomass in the defoliated birch seedlings, but not in the control ones. This tends to indicate a stronger carbon limitation of ectomycorrhizal colonization in defoliated seedlings.     

Plant responses and herbivory following simulated browsing and stem cutting of Combretum apiculatum

Abstract. Time and mode of herbivory on savanna trees and their subsequent responses are dependent on, among other things, earlier herbivory and fire. We used clipping (simulated browsing) and stem cutting (simulated heavy browsing and to some extent simulated fire) to evaluate such interactions. Study organisms were a deciduous, broad‐leaved tree species, Combretum apiculatum (Combretaceae), browsing large herbivores and leaf‐eating insects. The treatments were done in the late dry season before bud break. Late in the following wet season, we recorded plant responses to treatment and browsing. The treated trees, especially the cut ones, responded by producing larger and fewer annual shoots. Compared to control trees, there was a slight increase in shoot biomass of clipped trees and a strong reduction of cut ones. Leaf area increased in clipped trees, but decreased in cut ones. A marked increase in the number of browsed trees was recorded amongst treated trees. Number of bites, consumption and utilization also increased with severity of treatment. In contrast, insect herbivory was reduced on both clipped and cut trees. The observed patterns are discussed in relation to current ideas on plant ‐ herbivore interactions.     

Compensatory growth response of the legume, Medicago sativa, to defoliation and denodulation

A laboratory study was conducted to determine the effects of defoliation and denodulation on compensatory growth of Medicago sativa (L.). Plants grown hydroponically in clear plastic growth pouches were subjected to 0 and 50% nodule pruning, and 0, 25, 50, and 75% defoliation by clipping trifoliate leaves. An additional experiment was conducted to determine if clipping leaves simulated herbivory by Hypera postica (Gyllenhal) larvae. Previously, we determined that nodule pruning accurately simulated herbivory by Sitona hispidulus (L.) larvae (Quinn & Hall, 1992). Results indicated that denodulation stimulated nodule growth and caused exact compensation in standing and total number of nodules per plant within 15 days and in standing nodule biomass within 22 days of treatment. Denodulation caused a significant reduction (13%) in final shoot biomass, but did not affect significantly final root biomass. Percentage of change in number of trifoliate leaves per plant increased with the level of defoliation. Within 22 days of treatment, total number of trifoliate leaves per plant was similar to controls. However, final standing shoot biomasses were significantly less that controls, indicating undercompensatory growth. Shoot biomasses of the 25-, 50-, and 75%-defoliated plants were 18, 20, and 36% lower than controls, respectively. Nodule biomass per plant was reduced by 24 and 32% in 50- and 75%-defoliated plants, respectively, but was not affected significantly by 25% defoliation. Root biomass was affected by all levels of defoliation. Clipping trifoliate leaves accurately simulated defoliation by H. postica larvae. Our results indicated that partial defoliation affected shoot, root, and nodule biomass of M. sativa, but that partial denodulation only affected shoot biomass.     

不同生境下空心莲子草响应模拟昆虫采食的生长和化学防御策略

外来植物往往可以入侵多种生境并受到多种昆虫的采食,而不同生境条件将可能会影响这些入侵植物对昆虫采食的防御策略。以入侵我国的克隆植物——空心莲子草为研究对象,分别选择生长在水生生境、水陆两栖生境和陆生生境中的无性个体(分株),通过50%去叶处理模拟昆虫采食,分析不同生境下空心莲子草对模拟昆虫采食处理的生长及化学防御响应的差异。模拟昆虫采食处理显著抑制了陆生生境、水陆两栖生境以及水生生境下空心莲子草的根、茎、叶和总生物量,但对3种生境下空心莲子草的生物量分配(根冠比、根生物量分配、茎生物量分配和叶生物量分配)均无显著影响。陆生生境下空心莲子草根、茎和总生物量显著高于水陆两栖生境和水生生境,根冠比显著低于水陆两栖生境和水生生境。模拟昆虫采食处理显著降低了空心莲子草的木质素含量,而对单宁和总酚含量影响不显著。生境对木质素含量无显著影响,但陆生生境下空心莲子草单宁含量显著高于水陆两栖生境和水生生境,且总酚含量显著高于水陆两栖生境,表明陆生生境中空心莲子草具有更强的防御能力。空心莲子草木质素含量与总生物量无显著相关性,但在模拟采食情况下,其总酚含量与总生物量呈显著负相关,而无论模拟昆虫采食处理存在与否,空心莲子草单宁含量与总生物量均呈显著正相关。因此,空心莲子草存在昆虫介导的生长和化学防御之间的权衡,在昆虫采食的情况下可通过减少生长来增加对化学防御物质的投入,但生境对空心莲子草这种生长-防御权衡的影响十分有限。     

Growth responses of a woody species to clipping and goat saliva

Studies on the role of mammalian herbivore saliva in plant–animal interactions have mostly focused on graminoid species and bovine saliva. A trial was performed in Botswana with clipping treatments to simulate browsing of shoots and the application of goat saliva on the woody species Combretum apiculatum Sonder (Combretaceae). Treatments were performed during early growing season while shoot growth was rapid, and responses of trees were recorded later in the same season. Clipped shoots with saliva had significantly enhanced shoot growth (tripled in length) and leaf production (2.7 times more leaves) compared to clipped shoots without saliva. However, unclipped shoots still grew more than clipped shoots, with or without saliva treatment.     

Leaf dynamics and insect herbivory in a Eucalyptus camaldulensis forest under moisture stress

Abstract The influence of soil moisture content on leaf dynamics and insect herbivory was examined between September 1991 and March 1992 in a river red gum (Eucalyptus camaldulensis) forest in southern central New South Wales. Long-term observations of leaves were made in trees standing either within intermittently flooded waterways or at an average of 37. 5m from the edge of the waterways. The mean soil moisture content was significantly (P≤0.05) greater in the waterways than in the non-flooded areas. Trees in the higher soil moisture regime produced significantly larger basal area increments and increased canopy leaf area. This increase in canopy leaf area was achieved, in part, through a significant increase in leaf longevity and mean leaf size. Although a greater number of leaves was initiated and abscissed per shoot from the non-flooded trees, more leaves were collected from litter traps beneath the denser canopies of the flooded trees. Consumption of foliage by insects on the trees subjected to flooding compared to the non-flooded trees was not significantly different. However, the relative impact of insect herbivory was significantly greater on the non-flooded trees. Leaf chewing was the most common form of damage by insects, particularly Chryso-melidae and Curculionidae. No species was present in outbreak during this study. Leaf survival decreased as the per cent area eaten per leaf increased. In addition, irrespective of the level of herbivory, leaf abscission tended to be higher in E. camaldulensis under moisture deficit. The influence of soil moisture content on the balance between river red gum growth and insect herbivory is discussed.     

Induced defences in the neotropical tree Bauhinia brevipes (Vog.) to herbivory: effects of damage-induced changes on leaf quality and insect attack

Induced defences to herbivory are physical, nutritional, and allelochemical traits that change in plants following damage or stress, and that reduce the performance and/or preference of herbivores. The aim of this study was to verify the occurrence and effect of induced responses in Bauhinia brevipes (Vog.) (Leguminosae) which defend it against herbivores, through the manipulation of its leaves, and their effects on herbivore foraging behaviour. We selected 15 plants in the field, and three shoots per plant were subjected to one of three treatments: (1) damaged shoots (simulation of the main types of foliar herbivory and insect exclusion); (2) damaged control shoots (insect exclusion); and (3) control shoots (not manipulated). Water and nitrogen content, tannin concentration, levels of herbivory, and shoot growth rates were compared among treatments. Leaf quality varied among treatments. Damaged leaves showed higher tannin concentration, and lower water and nitrogen content compared to undamaged leaves. On the other hand, they experienced higher rates of herbivory than leaves on control shoots. Moreover, shoots that were experimentally induced showed a higher increase in final shoot length. These results suggest that simulated herbivory on B. brevipes reduced the nutritional quality of its leaves and increased the amount of secondary compounds, therefore altering insect herbivore attack and increasing shoot performance.     

Effects of warming climate on early-season carbon allocation and height growth of defoliated mountain birches

Tree carbohydrate reserves are usually compromised following insect outbreak, which results in a delay in leaf emergence and a reduction in growth, especially in cold environments. However, in recent times, severe defoliation of subarctic mountain birches (Betula pubescens ssp. czerepanovii) by the winter moth (Operophtera brumata) has not induced such responses. This may be the result of a warming climate stimulating plant primary metabolism. We examined if increasing thermal sum (sum of daily mean temperatures above +5 °C, d.d.) and complete foliage loss affected the concentrations of carbohydrates in sap, juvenile leaves, and fine roots of mountain birches in northern Finland and Norway. The sampling was conducted at the beginning of the growing season, two years after the insect outbreak. We also investigated the morphologic properties of mature leaves and the shoot growth of the trees. Our results showed that the carbohydrate concentrations in leaves and roots (averages 67.8 and 12.5 mg g^?1 DW, respectively) decreased in defoliated trees with increasing thermal sum (>400 d.d.), whereas the response in intact trees was the opposite. The carbohydrates in the sap were unaffected by defoliation or thermal sum accumulation. The leaf area of mature leaves and the height growth of long shoots were greater in trees at warmer sites, irrespective of defoliation. However, defoliation increased the leaf weight per area (SLW: specific leaf weight). We conclude that under warmer growing conditions, low early-season leaf and fine root carbohydrate concentrations of previously defoliated trees cannot be used as indicators of aboveground growth.     

Herbivore effects on developmental instability and fecundity of holm oaks

Plants are able to compensate for loss of tissue due to herbivores at a variety of spatial and temporal scales, masking detrimental effects of herbivory on plant fitness at these scales. The stressing effect of herbivory could also produce instability in the development of plant modules, and measures of such instability may reflect the fitness consequences of herbivory if instability is related to components of plant fitness. We analyse the relationships between herbivory, developmental instability and production of female flowers and fruits of holm oak Quercus ilex trees by means of herbivore removal experiments. Removal of leaf herbivores reduced herbivory rates at the tree level, but had no effect on mean production of female flowers or mature fruits, whereas herbivory tended to enhance flower production and had no effect on fruit abortion at the shoot level. Differences in herbivory levels between shoots of the same branch did not affect the size and fluctuating asymmetry of intact leaves. These results indicate compensation for herbivory at the tree level and over-compensation at the shoot level in terms of allocation of resources to female flower production. Removal of insect herbivores produced an increase in the mean developmental instability of leaves at the tree level in the year following the insecticide treatment, and there was a direct relationship between herbivory rates in the current year and leaf fluctuating asymmetry the following year irrespective of herbivore removal treatment. Finally, the production of pistillate flowers and fruits by trees was inversely related to the mean fluctuating asymmetry of leaves growing the same year. Leaf fluctuating asymmetry was thus an estimator of the stressing effects of herbivory on adult trees, an effect that was delayed to the following year. As leaf fluctuating asymmetry was also related to tree fecundity, asymmetry levels provided a sensitive measure of plant performance under conditions of compensatory responses to herbivory.     

Interactive effects of soil fertility and herbivory on Brassica nigra

Soil nutrient availability may affect both the amount of damage that plants receive from herbivores and the ability of plants to recover from herbivory, but these two factors are rarely considered together. In the experiment reported here, I examined how soil fertility influenced both the degree of defoliation and compensation for herbivory for Brassica nigra plants damaged by Pieris rapae caterpillars. Realistic levels of defoliation were obtained by placing caterpillars on potted host plants early in the life cycle and allowing them to feed until just before pupation on the designated plant. Percent defoliation was more than twice as great at low soil fertility compared to high (48.2% and 21.0%, respectively), even though plants grown at high soil fertility lost a greater absolute amount of leaf area (38.2 cm² and 22.1 cm², respectively). At both low and high soil fertility, total seed number and mean mass per seed of damaged plants were equivalent to those of undamaged plants. Thus soil fertility did not influence plant compensation in terms of maternal fitness. However, the pathways used to achieve compensation in seed production were different at low and high soil fertility. At low soil fertility, relative leaf growth rates (area added per inital area per day) of damaged plants were drastically reduced over the second week of caterpillar feeding. Damaged plants recovered the leaf area lost to herbivory in the two weeks following insect removal by increasing leaf relative growth rates above the levels seen for undamaged plants, but the replacement of leaf tissue lost to herbivory came at the expense of stem biomass. At high soil fertility, relative leaf growth rates of damaged plants were similar to those of undamaged plants both over the second week of caterpillar feeding and following caterpillar removal, and stem biomass was not affected by herbivory. These results suggest that higher levels of soil nutrients increased the ability of plants to stay ahead of their herbivores as they were being eaten. Because damaged plants at high soil fertility were able to maintain leaf growth rates to a greater extent than damaged plants at low soil fertility, they did not fall as far behind undamaged plants over the period of insect feeding and did not have as much catching up to do after feeding ended to compensate for herbivory.     

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.     

Growth responses of an African savanna tree, <Emphasis Type="Italic">Bauhinia thonningii</Emphasis> Schumacher,to defoliation,fire and climate

This study investigated the growth responses to defoliation, fire-exposure and climate factors of a widespread Africana savanna tree, Bauhinia thonningii Schumacher, at a site in central Zambia. Experimental trees (n = 47) were either exposed to fire (n = 12) in the first half of the dry season or protected from fire (n = 35). Some of the fire-protected trees (n = 12) were subjected to artificial defoliation in two consecutive years. Phenological responses (bud break, leaf flush and leaf production) to fire-exposure and defoliation were monitored on permanently marked sample shoots over a 2-year period. Radial tree growth (diameter at 1.3 m above ground) was measured annually over a 7-year period from 1998 to 2005. During the first two years, defoliation and fire-exposure advanced the onset of bud break and leaf flush but fire-exposed trees produced significantly less leaves than did trees protected from fire, probably because scorching caused more severe shoot die-back than is normal. Leaf production was also significantly affected by experimental treatments and their interaction with year. Although treatments had significant short-term effects on radial growth, previous-year growth significantly influenced current-year growth, thereby confirming the existence of autocorrelation in the time-series growth data of B. thonningii. The interaction between previous-year growth and climate factors explained a significant proportion (25–40%, P < 0.001) of the variance in annual tree growth. Variogram models predicted that a 2-year manual defoliation treatment would shorten the longer-term growth cycle while continuous fire-exposure extended the cycle by one year. The results are useful for the management of savanna trees.     

Effects of simulated herbivory in three old field Compositae with different inflorescence architectures

The effects of simulated herbivory (early or late defoliation and cutting of the flowering shoot) on the growth and reproduction of three species of monocarpic composite forbs (Crepis pulchra, Picris hieracioides and C. foetida) with different inflorescence architectures were studied in experimental plots. For the three species studied, early defoliation had no significant effect on subsequent growth. In contrast, late defoliation, occurring at the start of the season of drought, had a negative effect on growth and reproduction in the two Crepis species, particularly C. foetida, but had less effect on P. hieracioides. Sexual biomass was more clearly affected by late defoliation than was vegetative biomass, although the effects differed markedly among species possibly as a result of differences in phenology. Clipping the flowering shoot removed about 3 times less biomass than late defoliation and had little effect on vegetative biomass. It had much greater effects on the sexual biomass in P. hieracioides and C. pulchra, and resulted in the production of many shoots sprouting from the rosette, allowing the treated plants to regain a vegetative biomass close to that of control plants. Clipping did however lead to the production of shorter shoots and a reduction in the number of capitula formed. In C. foetida, much branching occurred even when the main shoot was not cut; the architecture of individual plants was therefore only slightly changed by clipping the apical bud and the sexual biomass of this species was not affected by ablation of the flowering shoot. Overcompensation was found in only two families of C. pulchra for vegetative biomass. No over-compensation was found for sexual biomass, despite an increase in the number of flowering shoots in C. pulchra and P. hieracioides following clipping. However situations close to compensation for the vegetative biomass in the three species and in P. hieracioides for the sexual biomass were recorded. The response of the three study species to simulated herbivory were related to their architecture and to the time of defoliation.     

Dieback of rural eucalypts: Response of foliar dietary quality and herbivory to defoliation

Rural dieback of Eucalyptus blakelyi trees growing on pastoral properties near Canberra is associated with chronic defoliation by insects. In order to test the effect of defoliation on subsequent herbivory, I artificially defoliated three healthy trees by clipping their terminal branchlets. The foliage that regrew on the clipped trees was nutritionally superior to the foliage it replaced, and was much more heavily damaged by grazing insects. There was a transient increase in the tannin content of the regrowth foliage, but this was apparently ineffective in defending it from subsequent herbivory. Compared with the foliage on nearby E. blakelyi trees that also produced major flushes of leaf growth during the same period, the regrowth on the clipped trees had enhanced dietary qualities and suffered more insect damage. Leaf age contributed to many of the differences in dietary quality, but when adjustments were made for the effects of leaf age the same trends remained. Five of the nearby trees were suffering from the chronic insect grazing associated with rural dieback, and the other five appeared healthy. The dietary quality of regrowth foliage on the clipped trees was qualitatively more similar to that of foliage on the dieback trees. Thus the chronic herbivory associated with rural dieback may be partly self-perpetuating, given this positive feedback between defoliation and dietary quality, and an apparent absence of other effective controls on insect populations.     

Compensation to simulated insect leaf herbivory in wild cotton (Gossypium hirsutum): responses to multiple levels of damage and associated traits

• Identifying the mechanisms of compensation to insect herbivory remains a major challenge in plant biology and evolutionary ecology. Most previous studies have addressed plant compensatory responses to one or two levels of insect herbivory, and the underlying traits mediating such responses remain elusive in many cases.
• We evaluated responses associated with compensation to multiple intensities of leaf damage (0% control, 10%, 25%, 50%, 75% of leaf area removed) by means of mechanical removal of foliar tissue and application of a caterpillar (Spodoptera exigua) oral secretions in 3‐month‐old wild cotton plants (Gossypium hirsutum). Four weeks post‐treatment, we measured plant growth and multiple traits associated with compensation, namely: changes in above‐ and belowground, biomass and the concentration of nutrients (nitrogen and phosphorus) and non‐structural carbon reserves (starch and soluble sugars) in roots, stems and leaves.
• We found that wild cotton fully compensated in terms of growth and biomass allocation when leaf damage was low (10%), whereas moderate (25%) to high leaf damage in some cases led to under‐compensation. Nonetheless, high levels of leaf removal (50% and 75%) in most cases did not cause further reductions in height and allocation to leaf and stem biomass relative to low and moderate damage. There were significant positive effects of leaf damage on P concentration in leaves and stems, but not roots, as well as a negative effect on soluble sugars in roots.
• These results indicate that wild cotton fully compensated for a low level of leaf damage but under‐compensated under moderate to high leaf damage, but can nonetheless sustain growth despite increasing losses to herbivory. Such responses were possibly mediated by a re‐allocation of carbohydrate reserves from roots to shoots.

     

Ecophysiological responses of salt cedar (Tamarix spp. L.) to the northern tamarisk beetle (Diorhabdacarinulata Desbrochers) in a controlled environment

The northern tamarisk beetle (Diorhabda carinulata Desbrochers) was released in several western states as a biocontrol agent to suppress Tamarix spp. L. which has invaded riparian ecosystems; however, effects of beetle herbivory on Tamarix physiology are largely undocumented and may have ecosystem ramifications. Herbivory by this insect produces discoloration of leaves and premature leaf drop in these ecosystems, yet the cause of premature leaf drop and the effects of this leaf drop are still unknown. Insect herbivory may change leaf photosynthesis and respiration and may affect a plant’s ability to regulate water loss and increase water stress. Premature leaf drop may affect plant tissue chemistry and belowground carbon allocation. We conducted a greenhouse experiment to understand how Tamarix responds physiologically to adult beetle and larvae herbivory and to determine the proximate cause of premature leaf drop. We hypothesized that plants experiencing beetle herbivory would have greater leaf and root respiration rates, greater photosynthesis, increased water stress, inefficient leaf nitrogen retranslocation, lower root biomass and lower total non-structural carbohydrates in roots. Insect herbivory reduced photosynthesis rates, minimally affected respiration rates, but significantly increased water loss during daytime and nighttime hours and this produced increased water stress. The proximate cause for premature leaf drop appears to be desiccation. Plants exposed to herbivory were inefficient in their retranslocation of nitrogen before premature leaf drop. Root biomass showed a decreasing trend in plants subjected to herbivory. Stress induced by herbivory may render these trees less competitive in future growing seasons.     

Seasonality of insect herbivory on the leaves of Neoboutonia macrocalyx in the Kibale National Park, Uganda

The seasonality of herbivory on the leaves of Neoboutonia macrocalyx Pax. in Kibale Forest National Park, Uganda, was studied. A total of 2929 fallen leaves was collected during 15 months under randomly-selected trees in three different habitats; natural forest and two selectively cut forest sites. The percentage of leaf area eaten and leaf size were estimated. Leaf herbivory was highly seasonal and correlated with rainfall in the previous 2 months, but less than 100 mm monthly rainfall had no effect. There was no correlation between leaf size and rainfall. Although Kibale Forest has two wet seasons, insect feeding on leaves had only one peak during the major rainy season from September to December. Three to four months after peak herbivory, leaves had very low rates of insect damage. Habitat had only a small effect on the amount of insect feeding. The sampling time accounted for 71% of variation in leaf herbivory. New leaves were formed continuously year-round. The constant leaf production by Neoboutonia trees may be an adaptation to escape generalist herbivorous insects which might be synchronized with the major wet season when the leaf flush of the most other deciduous species occurs. Thus, the availability of fresh leaves is not acting as a regulating factor in seasonality of Neoboutonia herbivory.     

PHOTOSYNTHESIS RESPONSE OF ‘CAROLINA’ CUCUMBER TO SIMULATED AND ACTUAL STRIPED CUCUMBER BEETLE (COLEOPTERA: CHRYSOMELI-DAE) DEFOLIATION*

Abstract Field studies were conducted in 1995 to compare the photosynthesis response between simulated striped cucumber beetle, Acalymma vittatum (F.), defoliation and actual A. vittatum defoliation in ‘Carolina’ cucumber. Six simulated defoliation levels (0%–100%)were used over 5 timings of defoliation (first true leaf to harvest). Plots were defoliated with scissors twice each week throughout each timing interval, and defined as continuous defoliation. In addition to continuous defoliation, one-time simulated defoliation treatments were imposed using the same timing treatments but only 4 defoliation levels (0, 25, 50, and 100%). Two cage studies, with A. vittatum adults, were used to provide estimates of actual insect defoliation injury. Beetles were placed in cages for 2 weeks at densities of 0, 1, 3, 5, and 10/plant and net photosynthetic rates were measured. No significant differences (P <0. ⁰⁵ were detected in pho-tosynthetic rates between simulated and actual insect defoliation treatments. Also, few significant differences in photosynthetic rates were detected between damaged and undamaged leaves in the simulated insect defoliation study. Results indicated that the simulated insect defoliation procedure appeared to accurately reflect the growth response of ‘Carolina’ cucumber subjected to actual A. vittatum defoliation, and that simulated defoliation results should be appropriate for developing economic injury levels for A. vittatum on cucumber.     

Consequences of the Elimination of Old Leaves upon Spring Phenological Events and the New Leaves Nutrient Concentration in a Wintergreen Woody Species in the Southern Hemisphere

Previous studies analyzed the importance of old leaves conservancy for wintergreen species plant growth only after early spring old leaves elimination. However, carbon and nutrient resources for growth could have already been translocated from old leaves to shoots during autumn. In this work, the effect of old leaves absence on the leaf mass per area (LMA, g m⁻²) and nutrient concentration of new spring leaves, shoot growth, and flowering was studied in Aristotelia chilensis, an Andean Patagonic woody wintergreen species of Argentina. Plants were studied after autumn defoliation (AD) or late winter defoliation (WD) and results were compared to those of undamaged control plants (CO). The new leaves LMA and mineral nutrient (N, P, K, and Mg) concentration values did not decrease in AD or WD compared to CO plants. Conversely, CO plants showed higher flowering intensity and shoot lengthening compared to AD or WD plants. There were not remarkable differences regarding the defoliation time, though non-flowering shoots grew in a lesser degree than the flowering shoots in WD plants. It was concluded that A. chilensis old leaves cohort is an important source to shoot growth and flowering but their absence does not affect the new leaves structure or nutritional status from early spring in either AD or in WD plants. New leaves formation probably is guaranteed by resources (carbon and nutrients) previously stored in stems or even in the buds containing the preformed leaves since March, by the end of summer. Provided the availability of complete resources for the new leaf flush independently of the old leaves A. chilensis would restore the carbon balance as soon as possible to resume the growth of heterotrophic tissues at normal rates. Endogenous response to counterbalance the old leaves absence on non-flowering shoots was more effective when there was greater lag time between defoliation and shoot growth resume. Flowering and non-flowering shoots compete for the available resources when A. chilensis have not yet expanded leaves and shoots supporting reproductive structures were stronger sinks compared to non-flowering shoots in WD plants.