High abscission rates of damaged expanding leaves: field evidence from seedlings of a Bornean rain forest tree

Herbivore damage is known to cause the premature loss of mature leaves. However, the effects of herbivory on abscission during the early stages of leaf development remain unexplored, even though herbivores frequently prefer unlignified, immature leaves. In a field experiment, we removed 50% of the tissue from leaves at various stages of development on seedlings of Shorea hopeifolia (Dipterocarpaceae), a dominant rain forest tree in Indonesian Borneo. Four weeks following simulated herbivory, >88% of unlignified expanding leaves had been abscised, compared to only 20% of fully expanded, unlignified leaves and 0% of recently lignified, mature leaves. In a separate experiment over 9 wk, simulated herbivory did not increase abscission rates of mature leaves, even when 75% of leaf tissue was removed. Because most (58%) of S. hopeifolia seedlings under natural conditions had lost 1% or less of the tissue from their mature leaves, herbivore damage probably has little effect on the abscission of mature leaves. In contrast, the tendency for damaged expanding leaves to abscise may explain why 49% of S. hopeifolia seedlings had already lost their youngest leaf. If similar patterns occur in other species, herbivore attacks on developing leaves may contribute substantially to both leaf loss and the cumulative impact of herbivory on the growth and survival of whole plants.     

Simulated herbivory advances autumn phenology in Acer rubrum

To determine the degree to which herbivory contributes to phenotypic variation in autumn phenology for deciduous trees, red maple (Acer rubrum) branches were subjected to low and high levels of simulated herbivory and surveyed at the end of the season to assess abscission and degree of autumn coloration. Overall, branches with simulated herbivory abscised ～7 % more leaves at each autumn survey date than did control branches within trees. While branches subjected to high levels of damage showed advanced phenology, abscission rates did not differ from those of undamaged branches within trees because heavy damage induced earlier leaf loss on adjacent branch nodes in this treatment. Damaged branches had greater proportions of leaf area colored than undamaged branches within trees, having twice the amount of leaf area colored at the onset of autumn and having ～16 % greater leaf area colored in late October when nearly all leaves were colored. When senescence was scored as the percent of all leaves abscised and/or colored, branches in both treatments reached peak senescence earlier than did control branches within trees: dates of 50 % senescence occurred 2.5 days earlier for low herbivory branches and 9.7 days earlier for branches with high levels of simulated damage. These advanced rates are of the same time length as reported delays in autumn senescence and advances in spring onset due to climate warming. Thus, results suggest that should insect damage increase as a consequence of climate change, it may offset a lengthening of leaf life spans in some tree species.     

Precipitation on abscised leaves modulates leaf‐mining beetle (Trachys yanoi) survivorship and population dynamics

1. We clarified the effects of early leaf abscission on the survivorship of the leaf‐mining beetle Trachys yanoi Y. Kurosawa (Coleoptera: Buprestidae) and the underlying mechanism in relation to weather conditions in Japan. Trachys yanoi is an insect pest of zelkova trees [Zelkova serrata (Thunb.) Makino (Rosales: Ulmaceae)]. Larvae burrow into zelkova leaves and feed on leaf tissue, causing early leaf abscission. 2. This study investigated the relationship between the beetle population and weather conditions over 10 years in a zelkova forest. The effects of moisture and temperature on adult emergence from early abscised leaves were examined in the laboratory and in the field. 3. The beetle population in the studied forest was negatively affected by high precipitation levels when the beetles still inhabited early abscised leaves. Fewer adults emerged from early abscised leaves under wet conditions than under dry conditions, in both laboratory and field tests. 4. These results demonstrate that early leaf abscission plays an important role in leaf‐mining beetle survivorship and population dynamics, and that the amount of precipitation when leaf‐mining beetles still inhabit early abscised leaves modulates this effect. 5. Because precipitation when leaf‐mining beetles still inhabit early abscised leaves was mainly driven by an East Asian rainy season front, the beetle population dynamics were affected by the activity of the front.     

Leaf abscission phenology of a scrub oak: consequences for growth and survivorship of a leaf mining beetle

Brachys tessellatus is a leaf-mining beetle that attacks Quercus laevis (turkey oak), a deciduous scrub oak in the fall line Sandhills of the southeastern United States. This oak species varies substantially in leaf abscission phenology. In the fall of 1994 we examined leaf abscission patterns at three sites in central South Carolina and found that leaves containing active miners abscised earlier than leaves without active miners. To quantify the effect of leaf abscission phenology on the survivorship and body size of leaf miners within the abscised leaves, we marked naturally and manually abscised leaves, left leaves on the ground to over-winter, and then collected leaves before beetles emerged the following spring. We found that beetles inside leaves that were abscised early in the season were more likely to be killed by predators, and that they produced smaller pupae than beetles within leaves abscised later in the season. We also evaluated the relationship between adult beetle body size and fecundity; body size of B. tessellatus is positively correlated with lifetime fecundity and daily oviposition rate indicating that effects of leaf-abscission phenology on beetle body size likely translate into effects on beetle fitness. We discuss how B. tessellatus has adapted to variation of leaf abscission phenology of oaks by evolving developmental plasticity that permits successful completion of larval development over a range of season lengths.     

Exposure of cottonwood plants to ozone alters subsequent leaf decomposition

Summary Cottonwood saplings were exposed to ozone or charcoal-filtered air in a closed chamber. After leaf abscission, decomposition of individual leaf discs was measured in containers of stream water. Exposure of plants to 200 ppb ozone for 5 h caused early leaf abscission and changes in the chemical composition of leaves at time of abscission. Early-abscised leaves from O₃-exposed plants had higher nitrogen, but decomposed more slowly than leaves from control plants. Leaves from O₃-exposed plants that abscised at the normal time had lower nitrogen content and lower specific leaf mass than control leaves, but decomposed at the same rate as leaves from control plants. The results imply that O₃ exposure can alter fundamental processes important to the functioning of detritus-based aquatic ecosystems.     

Ethylene-induced leaf abscission in cotton seedlings : the physiological bases for age-dependent differences in sensitivity

The speed of ethylene-induced leaf abscission in cotton (Gossypium hirsutum L. cv LG-102) seedlings is dependent on leaf position (i.e. physiological age). Fumigation of intact seedlings for 18 hours with 10 microliters per liter of ethylene resulted in 40% abscission of the still-expanding third true (3°) leaves but had no effect on the fully expanded first true (1°) leaves. After 42 hours of fumigation with 50 microliters per liter of ethylene, total abscission of the 3° leaves occurred while <50% abscission of the 1° leaves was observed. On a leaf basis, endogenous levels of free IAA in 1° leaves were approximately twice those of 3° leaves. Free IAA levels were reduced equally (approximately 55%) in both leaf types after 18 hours of ethylene (10 microliters per liter) treatment. Ethylene treatment of intact seedlings inhibited the basipetal movement of [¹⁴C]IAA in petiole segments isolated from both leaf types in a dose-dependent manner. The auxin transport inhibitor N-1-naphthylphthalamic acid increased the rate and extent of ethylene-induced leaf abscission at both leaf positions but did not alter the relative pattern of abscission. Abscission-zone explants prepared from 3° leaves abscised faster than 1° leaf explants when exposed to ethylene. Ethyleneinduced abscission of 3° explants was not appreciably inhibited by exogenous IAA while 1° explants exhibited a pronounced and protracted inhibition. The synthetic auxins 2,4-D and 1-naphthaleneacetic acid completely inhibited ethylene-induced abscission of both 1° and 3° explants for 40 hours. It is proposed that the differential abscission response of cotton seedling leaves is primarily a result of the limited abscission-inhibiting effects of IAA in the abscission zone of the younger leaves.     

Inhibition of premature leaf abscission by a leafminer and its adaptive significance

We tested the possibility that a lepidopteran leafminer, Coptotriche japoniella Puplesis and Diskus, inhibits the host plant Eurya japonica Thunberg from abscising mined leaves prematurely to increase its survivorship in immature stage. We monitored abscission patterns of mined leaves with sacrificed larvae, mined leaves with living larvae, and unmined leaves from April to July 2004 and 2005 until leafminers emerged as adults. Unmined leaves rarely abscised before July. Mined leaves with sacrificed larvae fell at a constant rate after May, abscising significantly more than unmined leaves. In contrast, mined leaves with living larvae rarely fell before adult emergence; afterward they abscised rapidly. We also examined larval/pupal survivorship and mortality sources on the ground and trees after leafminers completed larval development. Leafminers on the ground suffered a higher mortality from predation than those on trees, and thus they emerged as adults on the ground less successfully. These findings suggest that the leafminer C. japoniella prevents the host plant from abscising mined leaves prematurely until adult emergence, thereby increasing their survivorship.     

Herbivory mitigation through increased water-use efficiency in a leaf-mining moth-apple tree relationship

Herbivory alters plant gas exchange but the effects depend on the type of leaf damage. In contrast to ectophagous insects, leaf miners, by living inside the leaf tissues, do not affect the integrity of the leaf surface. Thus, the effect of leaf miners on CO2 uptake and water-use efficiency by leaves remains unclear. We explored the impacts of the leaf-mining moth Phyllonorycter blancardella (Lepidoptera: Gracillariidae) on light responses of the apple leaf gas exchanges to determine the balance between the negative effects of reduced photosynthesis and potential positive impacts of increased water-use efficiency (WUE). Gas exchange in intact and mined leaf tissues was measured using an infrared gas analyser. The maximal assimilation rate was slightly reduced but the light response of net photosynthesis was not affected in mined leaf tissues. The transpiration rate was far more affected than the assimilation rate in the mine integument as a result of stomatal closure from moderate to high irradiance level. The WUE was about 200% higher in the mined leaf tissues than in intact leaf portions. Our results illustrate a novel mechanism by which plants might minimize losses from herbivore attacks; via trade-offs between the negative impacts on photosynthesis and the positive effects of increased WUE.     

Seasonal patterns of herbivory,leaf traits and productivity consumption in dry and wet Patagonian forests

1. Endemic herbivory can influence forest ecosystem function, but how annual productivity consumption relates to seasonal resource utilisation by folivore guilds remains poorly understood. 2. Monthly changes in leaf damage and foliage traits were monitored in ‘dry’ and ‘wet’Nothofagus pumilio (Fagales: Nothofagaceae) deciduous forests in northern Patagonia, Argentina. Herbivore‐induced leaf abscission was assessed and foliar productivity consumption was measured in the canopy and in litterfall harvests. 3. Seasonal damage ranged from 8% to 32% in dry forest, but remained below 5% in wet forest although foliar quality was higher in the latter. In dry forest, dominant guilds were temporally separated; leaf miners consumed younger foliage in spring to early summer, whereas leaf tiers prevailed in late summer to autumn. In wet forest, damage created by external chewers was concentrated in early summer. 4. Insect damage induced premature leaf abscission, especially in dry forest. Although foliar production in wet forest doubled that in dry forest, the percentage of productivity lost to folivores was higher in dry (14–20%) than in wet (1.2–1.8%) forest. 5. The overall greater impact of herbivory in dry forest canopies countered the expectation that consumption would increase with plant productivity and nutritional quality. Lower temperatures and a shorter growing season are likely to constrain folivory in wet forest stands.     

Delayed greening,leaf expansion,and damage to sympatric<Emphasis Type="Italic"> Shorea</Emphasis> species in a lowland rain forest

The function of delayed greening in the seedlings of canopy tree species in a lowland tropical rain forest was examined in terms of its potential defensive value against herbivory. To explore the ecological and evolutionary backgrounds for delayed greening, we chose eight sympatric congeneric (Shorea) dipterocarp species that were either normal-greening or delayed-greening species. Expansion and toughening of leaves took approximately 30 days for all species, and did not differ between the normal- and delayed-greening species. The main factors that affected leaf damage during expansion were insect herbivory and fungal infection. Levels of leaf damage were significantly lower for delayed-greening species than for normal-greening species, but proportions of heavily damaged leaves and leaf abscission during expansion did not differ. In addition, no significant difference was found in damage levels on leaves (aged 1–2 months) of naturally occurring seedlings between normal- and delayed-greening species. Therefore, delayed greening may effectively reduce the level of leaf damage in young expanding leaves, but may not necessarily reduce leaf abscission and damage to mature leaves. The existence of delayed greening could not be simply explained by the phylogenetic and ecological backgrounds of the trees. Consequently, delayed greening may have a function in reducing damage during expansion, but more information (such as knowledge of the secondary metabolites involved in this phenomenon) is needed to explain fully why these species exhibit delayed greening.     

Factors underlying the late seasonal appearance of the lepidopterous leaf-mining guild on oak

Abstract. 1. Most lepidopterous leaf mining species found on the oak Quercus robur in Britain develop in summer. At this time of year, externally feeding caterpillars remove little leaf area since most of these free living folivores are predominantly spring feeders.
2. I forced Phyllonorycter harrisella (L.) miners to oviposit in spring, then exposed developing larvae to a wide range of leaf damage levels.
3. Leaf miner survivorship and mean female pupal weight were significantly greater in the experimental spring generation on undamaged oak leaves, this being when oak foliage is of the highest nutritional quality.
4. Leaf miner survivorship in all generations is correlated with natural leaf damage levels. Experimental leaf damage also reduces miner survivorship.
5. Leaf damage reduces miner survivorship by increasing the probability of larval death due to wound induced responses.
6. The role of asymmetrical competitive interactions between caterpillars and leaf-miners in determining the late seasonal appearance of miners is discussed. It is suggested that wound induced responses may play an important part in the structuring of phytophagous insect communities.     

The distribution pattern of herbivory in a beech canopy

Abstract. (1) The spatial distribution pattern of herbivory by phyllophagous canopy arthropods in a 90-year-old beech stand was analysed.
(2) Two main patterns of spatial distribution were recorded. The leaf mining moths Phyllonorycter maestingella and Stigmella hemargyrella and the adult weevils Rhynchaenus fagi and Phyllobius argentatus mainly attacked the under-story and the latter species especially the low canopy in the central part of the stand. Mines of R. fagi chiefly occurred in the top of the overstory canopy or at the fringe of the stand. However, comparable distribution patterns are only expected in mature beech stands composed of distinct overstory and understory layers.
(3) The effect of the actual spatial distribution pattern of herbivory on photosynthesis, productivity, and transpiration of beech in the research stand is discussed. It is suggested that in some cases the shade leaves of the understory canopy may cushion the effect of herbivory in the beech stand.     

The deleterious effects of salinity stress on leafminers and their freshwater host

Abstract.  1. Salinity is an important cause of abiotic stress in wetland communities yet little is known about its consequences for freshwater plants and their insect herbivores. The goal of this study was to test the effect of salinity on a leafmining insect, Cerodontha iridiphora , and its herbaceous host plant, Iris hexagona .
2. Leafminer performance was evaluated on irises grown in control and saline treatments, and the effects of salinity and herbivory on leaf quality and mortality was measured.
3. Leafminer density and size were significantly lower on irises grown in saline water compared with freshwater.
4. Both salinity and herbivory accelerated leaf senescence and mortality, and their combined effects increased tissue loss by an order of magnitude compared with controls.
5. Leafminers acted as nutrient sinks. The undamaged regions of mined leaves contained 40% less nitrogen than unmined leaves, providing a mechanism for the premature leaf mortality.
6. Salinity was detrimental to the performance and survival of both the iris leafminer and its host plant. We propose that glycophytic host plants and their insect herbivores will suffer more than halophytic communities from environmental salinity because they lack the adaptive mechanisms to tolerate this potent physiological stress.     

Oviposition site preference and larval mortality in a leaf-mining moth

Abstract. 1. The univoltine leaf-mining moth, Lithocolletis quercus Ams., is endemic to Israel, where it spends its 10.5 month larval period feeding only in the leaves of Quercus calliprinos Webb.
2. We compared patterns of egg deposition and sources of larval mortality to test whether oviposition patterns and site preferences confer an enhanced likelihood of larval survival.
3. Dominant sources of larval mortality were premature leaf abscission and death from unknown causes, whereas predation, parasitism and intraspecific interference accounted for relatively little larval mortality.
4. Eggs, and thus mines, were aggregated among leaves of host trees even though premature leaf abscission was positively correlated with density of mines per leaf. Interference competition among larvae was the only other density-dependent mortality factor.
5. Oviposition patterns within leaves mitigated the probability of death from larval interference, and probably also from early leaf abscission.
6. Despite these density-dependent mortality factors, overall probability of larval survival to pupation was independent of initial density of mines on a leaf.
7. The long larval period allows synchrony between oviposition flights and times of predictable resource availability.     

伤害和挥发物对棉花花外蜜的诱导效应

植物花外蜜的分泌是一种植物间接防御反应。为了明确植食性昆虫、机械伤和机械伤诱导的挥发性气体在植物花外蜜诱导分泌中的作用,分析了咀嚼式口器昆虫棉铃虫Helicoverpa armigera(Hübner)、刺吸式口器昆虫棉蚜Aphis gossypii Glover取食、剪刀机械伤、剪刀机械伤+棉铃虫反吐物、针刺机械伤以及机械伤诱导挥发物、顺式-茉莉酮对棉花Gossypium hirsutum L.叶片花外蜜分泌量的影响。结果表明,棉铃虫取食、剪刀机械伤、剪刀机械伤+棉铃虫反吐物处理均显著增加了被处理叶片花外蜜的分泌量。棉花花外蜜的诱导效应在处理叶片上表现明显,并且在较幼嫩的第3片真叶上也有系统性增长。顺式-茉莉酮和机械伤挥发物处理1 d对棉花较幼嫩的第4、5片真叶花外蜜有诱导效应。棉花叶片花外蜜的诱导主要与植物组织损伤有关;不同口器类型的昆虫对棉花叶片花外蜜的诱导量不同,咀嚼式口器的棉铃虫对棉花花外蜜的诱导强度显著高于刺吸式口器的棉蚜;顺式-茉莉酮和机械伤诱导的挥发物能作为棉花植株间交流的信息物质诱导棉花幼嫩叶片花外蜜的分泌。     

Synergistic interactions between volicitin,jasmonic acid and ethylene mediate insect-induced volatile emission in Zea mays

Plants display differential responses following mechanical damage and insect herbivory. Both caterpillar attack and the application of caterpillar oral secretions (OS) to wounded leaves stimulates volatile emission above mechanical damage alone. Volicitin ( N- 17-hydroxylinolenoyl- l -glutamine), present in beet armyworm (BAW, Spodoptera exigua ) OS, is a powerful elicitor of volatiles in excised maize seedlings ( Zea mays cv. Delprim). We consider some of the mechanistic differences between wounding and insect herbivory in maize by examining the activity of volicitin, changes in jasmonic acid (JA) levels, and volatile emission from both intact plant and excised leaf bioassays. Compared to mechanical damage alone, volicitin stimulated increases in both JA levels and sesquiterpene volatiles when applied to intact plants. In a bioassay comparison, excised leaves were more sensitive and produced far greater volatile responses than intact plants following applications of both volicitin and JA. In the excised leaf bioassay, volicitin applications (10–500 pmol) to wounded leaves resulted in dose dependent JA increases and a direct positive relationship between JA and sesquiterpene volatile emission. Interestingly, volicitin-induced JA levels did not differ between intact and excised bioassays, suggesting a possible interaction of JA with other regulatory signals in excised plants. In addition to JA, insect herbivory is known to stimulate the production of ethylene. Significant increases in ethylene were induced only by BAW herbivory and not by either wounding or volicitin treatments. Using intact plant bioassays, ethylene (at 1 µl l⁻¹ or less) greatly promoted volatile emission induced by volicitin and JA but not mechanical damage alone. For intact plants, wounding, elicitor-induced JA and insect-induced ethylene appear to be important interacting components in the stimulation of insect-induced volatile emission.     

Effect of exposure to subfreezing temperatures on ethylene evolution and leaf abscission in citrus

Citrus leaves exposed to subfreezing temperatures evolved ethylene at rates between 0.1 and 38.3 microliters per kilogram fresh weight per hour whereas untreated leaves evolved between 0.01 and 0.50 microliter per kilogram fresh weight per hour. Leaves not injured by freezing temperatures did not abscise, and ethylene evolution was near normal after 2 days. Freeze-injured leaves continued evolving high ethylene levels 4 or 5 days subsequent to freeze injury, and many of the freeze-killed leaves abscised. Supportive evidence suggested freeze-induced ethylene was involved in freeze-induced leaf abscission; whereas freeze-inhibited abscission was not due to a lack of ethylene but injury to other metabolic systems necessary for abscission.     

Herbivory-induced stress: Leaf developmental instability is caused by herbivore damage in early stages of leaf development

Herbivory is a major source of plant stress and its effects can be severe, decreasing plant fitness, or subtle, affecting the development of leaves by influencing the normal pattern of growth and expansion of leaf blades. Fluctuating asymmetry (FA) analysis is recognized as a measure of plant stress, and can be used to evaluate subtle effects of herbivory on the imperfect growth of bilaterally symmetrical traits, such as leaves. One general issue is that authors usually consider FA as an indicator of stress, which can attract herbivores (plant stress hypothesis), and studies showing that herbivores themselves affect leaf symmetry (herbivory-induced stress hypothesis) are scarce, with mixed results. Here, we investigated the relationship between herbivory by thrips and leaf FA in Banisteriopsis malifolia and Heteropterys escallonifolia (Malpighiaceae). Pseudophilothrips obscuricornis is a free-living, non-pest, sucking species that feeds mainly on leaf buds. We hypothesized that herbivory by thrips in the early stages of leaf development would provoke increased FA levels in mature leaves. The results showed that thrips herbivory rate was low, affecting barely more than 1% of the leaf blade. Nonetheless, thrips-attacked leaves of B. malifolia and H. escallonifolia presented increases of 15 and 27% in leaf asymmetry, respectively, compared to uninjured leaves, corroborating the herbivory-induced stress hypothesis. Since herbivory by thrips in leaf buds was related to significant increases in the stress of mature leaves, we assume that under these circumstances, FA can be used as a biomarker for plant stress following herbivory damage. To be useful as a biomarker of stress, FA in plants must be investigated with caution, taking into account the natural history of the herbivore species and timing of leaf damage.     

The alkaloidal responses of wild tobacco to real and simulated herbivory

Summary I compared the induced alkaloidal response in undamaged leaves of plants subjected to herbivory by the larvae of Manduca sexta and to different simulations of this herbivory; all herbivory treatments removed similar amounts of leaf mass. Although larval feeding induced a significant increase (2.2x) in alkaloid concentrations compared to undamaged plants, the alkaloid responses to larval feeding were significantly lower than the responses to an herbivory simulation (4x controls) which involved removing the same amount of leaf area from the same positions on the leaf, over a similar time period. Moreover, another herbivory simulation, identical in amount of leaf mass removed and duration of damage to the larval feeding, but without regard to spatial array of leaf damage, resulted in an alkaloidal response (5.5x controls) higher still than the previous herbivory simulation. In a second experiment the importance of leaf vein damage on the induced alkaloidal response was examined. Here, leaf removal that involved cutting leaf tissues from between secondary veins before removing the midrib, resulted in alkaloidal responses that were significantly lower (1.7x controls) than responses from leaf removal that involved cutting both veins and midribs along with the intervein tissues (2.6x controls). Vein damage alone did not produce a significant response. These results indicate that herbivory is difficult to simulate: that how a leaf is damaged can be as important as the magnitude of leaf damage in determining a plant's response to damage.