Defoliation of the annual herb Abutilon theophrasti: mechanisms underlying reproductive compensation

A number of studies have shown that under some conditions plants may fully or partially compensate for leaf tissue loss; however, the mechanisms underlying compensatory responses are not well understood. Previous work demonstrated that the annual herb Abutilon theophrasti fully compensated for 75% defoliation, but only when grown in the absence of stem competition. We examined potential mechanisms of compensatory response and how they are influenced by resource limitation (i.e., competition for light). Full compensation for these annual plants was defined as equal final reproductive output in defoliated and control plants. In the current study we observed substantial compensation in defoliated plants growing at low density, despite losing 75% of leaf area prior to the onset of flowering. Plant responses associated with compensation included (1) increased reproductive efficiency, which may in turn may have resulted from increased canopy light penetration and transient increases in leaf-level photosynthetic rates; (2) greater allocation to reproduction (RA); (3) changes in biomass allocation from roots to shoots; (4) lower leaf longevity, and (5) increased percent fruit set. Although some of these responses were also observed in defoliated plants grown at high density, the inability of high-density plants to compensate appeared to result from competition for light; these plants delayed reproduction and continued to produce new leaves late in the growing season after low-density, defoliated plants had shifted allocation of resources to reproduction. Received: 20 June 1996 / Accepted: 12 February 1997     

Compensatory growth responses of seedlings of Pharbitis purpurea (Convulvulaceae) to tissue removal at different seeding depths of seeds

Tissue damage to seedlings can limit their later growth, and the further effects may be greater with increasing seedling age. Seedlings, however, can minimize the effect of damage through compensatory growth. Seedlings of Pharbitis purpurea grow in frequently disturbed habitats and generally tolerate damage to leaf tissues. We evaluated the compensatory responses of the cotyledon to different levels of defoliation and their effect on seedling growth and development. We also examined the relationship between seeding depth and compensatory growth. We tested seven defoliation treatments with one or both cotyledons and/or the apical meristem of seedlings removed from seeds buried at a seeding depth of either 2 or 5?cm. We then measured 12 growth traits of the seedlings to assess development and growth compensation. The area, thickness, biomass, and longevity of the remaining cotyledon were also measured to quantify increased growth as result of treatment effects at both seeding depths. The results showed that defoliation reduced seedling height, belowground length, and total biomass significantly in subsequent growth in all treatments. However, defoliation treatments had direct positive impacts on growth at 2?cm depth compared with 5?cm depth. In contrast, the compensation of cotyledon area (C _area), biomass (C _mass), and thickness (C _thickness) was greater at 5?cm depth than at 2?cm depth. The results thus indicate that P. purpurea seedlings adopted a compensatory growth strategy to resist leaf loss and minimize any adverse effects using the remaining cotyledon. Increasing seeding depth can aggravate the compensatory growth of remain cotyledon after partial defoliation.     

Overall grazing tolerance index (overall GTI) is not an ideal predictor for describing a single‐species tolerance to grazing

Plants' pattern of compensatory growth is often used to intuitively estimate their grazing tolerance. However, this tolerance is sometimes measured by the overall grazing tolerance index (overall GTI), which assumes that tolerance is a multivariate linear function of various underlying mechanisms. Because the interaction among mechanisms is not independent, the grazing tolerance expression based on overall GTI may be inconsistent with that based on compensatory growth. Through a manipulative field experiment from 2007 to 2012, we measured the responses of 12 traits of Elymus nutans to clipping under different resource availabilities in an alpine meadow and explored the compensatory aboveground biomass and the overall GTI to assess the possible differences between the two expressions of tolerance. Our results showed that these two expressions of tolerance were completely opposite. The expression based on overall GTI was over‐compensatory and did not vary with clipping and resource availability, while the expression based on compensatory aboveground biomass was under‐compensatory and altered to over‐compensation after fertilization. The over‐expression of highly variable traits with extremely high negative mean GTI to defoliation damage, the influence of random errors contained in traits considered, and the doubling weight of functional redundant traits greatly inflated the overall GTI, which leads to the inconsistency of the two tolerance expressions. This inconsistency is also associated with the different determining mechanisms of the two tolerance expressions. Our data suggest that plants' grazing tolerance is not a multivariate linear function of traits or mechanisms that determine grazing tolerance; the overall GTI is only a measure of traits' variability to defoliation damage. Our findings highlight that the tolerance of E. nutans mainly depends on the response of traits with lower variability to defoliation, and the overall GTI is not an ideal predictor for describing a single‐species tolerance to grazing.     

Growth,morphology and leaf characteristics after simulated herbivory in Chinese subtropical evergreen saplings

Growth, morphology and leaf characteristics were assessed in late spring following simulated autumnal defoliation in second-year saplings of three Chinese subtropical evergreen tree species.Castanopsis fargesii showed strong compensatory growth in terms of plant biomass after removal of both 50 and 75% of leaf biomass and slight compensatory growth after 90% defoliation. DefoliatedC. fargesii saplings had more leaves per unit shoot length than non-defoliated saplings. New leaves on defoliated plants were smaller and had higher per area nitrogen content than new leaves on non-defoliated plants.Pinus massoniana andElaeocarpus japonicus showed strong and no compensatory growth, respectively, after 50% defoliation. The strong compensatory growth inP. massoniana andC. fargesii may partly explain why these species predominate in the early and late successional phases of evergreen broad-leaved forests     

Production and nitrogen responses of the African dwarf shrub Indigofera spinosa to defoliation and water limitation

Summary The dwarf shrub Indigofera spinosa Forsk. (Papilionacea), a native forage species of arid Northwest Kenya, was propogated from seed, grown in a controlled environment, and subjected to three treatments of defoliation and watering frequencies in a factorial experimental design. Biomass production and nitrogen accumulation in tissue components were measured to determine defoliation responses in a water-limited environment. We hypothesized that plants would maintain biomass and nitrogen flows despite removal of aboveground meristems and tissues by defoliation. Principal experimental results included a slight reduction (11%; P=0.08) of total biomass production by clipping ca. 1/3 or 2/3 of new leaves and stems and all apical meristems every month. Total aboveground production was not affected by clipping, while final root biomass was reduced 17% by the 2/3 clipping. The least water stressed plants were affected most negatively by defoliation, and the unclipped plants responded more negatively to greater water limitation. Plants achieved partial biomass compensation through alterations in shoot activity and continued allocation of photosynthate to roots. A smaller fraction of leaf production was directed to litter in clipped plants although clipping only removed the youngest tissues, suggesting that clipping increased leaf longevity. In turn, each leaf probably contributed a greater total quantity of photosynthate. Photosynthetic rates were also likely to have been increased by clipping water-stressed plants. In contrast to biomass, plants overcompensated for nitrogen lost to defoliation. Total nitrogen uptake by individual plants was stimulated by defoliation, as there was more total nitrogen in leaves and stems. Increased nitrogen uptake was achieved by clipping stimulation of total uptake per unit of root rather than of total root mass.     

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.

     

Counteractive biomass allocation responses to drought and damage in the perennial herb Convolvulus demissus

Herbivory and water shortage are key ecological factors affecting plant performance. While plant compensatory responses to herbivory include reallocation of biomass from below‐ground to above‐ground structures, plant responses to reduced soil moisture involve increased biomass allocation to roots and a reduction in the number and size of leaves. In a greenhouse study we evaluated the effects of experimental drought and leaf damage on biomass allocation in Convolvulus demissus (Convolvulaceae), a perennial herb distributed in central Chile, where it experiences summer drought typical of Mediterranean ecosystems and defoliation by leaf beetles and livestock. The number of leaves and internode length were unaffected by the experimental treatments. The rest of plant traits showed interaction of effects. We detected that drought counteracted some plant responses to damage. Thus, only in the control watering environment was it observed that damaged plants produced more stems, even after correcting for main stem length (index of architecture). In the cases of shoot : root ratio, relative shoot biomass and relative root biomass we found that the damage treatment counteracted plant responses to drought. Thus, while undamaged plants under water shortage showed a significant increase in root relative biomass and a significant reduction in both shoot : root ratio and relative shoot biomass, none of these responses to drought was observed in damaged plants. Total plant biomass increased in response to simulated herbivory, apparently due to greater shoot size, and in response to drought, presumably due to greater root size. However, damaged plants under experimental drought had the same total biomass as control plants. Overall, our results showed counteractive biomass allocation responses to drought and damage in C. demissus. Further research must address the fitness consequences under field conditions of the patterns found. This would be of particular importance because both current and expected climatic trends for central Chile indicate increased aridity.     

Interactive effects of clipping and nutrient availability on the compensatory growth of a grass species

Resource availability is an important factor affecting the capacity of compensatory growth after grazing. We performed a greenhouse experiment with Poa bulbosa, a small perennial grass of the Mediterranean and Central Asian grasslands, to test the importance of nutrient availability for compensatory growth after clipping. We also compared the results with predictions of the limited resource model (LRM). Plants were grown at low and high fertilization levels and subjected to a clipping treatment. Contrary to the LMR, we found that in Poa plants compensatory growth occurred under the high fertilization level, while it did not occur under the low level. The LMR predicts a higher tolerance for grazing in the stressful environment. Our plants showed a significant decrease in their relative growth rates (RGR) after clipping. Although the plants allocated a 32–188% greater fraction of the mass to lamina growth after clipping, this greater allocation to the leaves did not fully compensate for the initial reduction in leaf area ratio (LAR). A sensitivity analysis showed for the clipped plants under the high fertilization treatment, that changes in leaf allocation (f _lam) enabled the plants to compensate for a part of the potential loss caused by defoliation. Probably, the increased biomass allocation comes largely from the bulbs. We conclude that the inconsistency of the LRM with our results originates in the lack of compensatory mechanisms in the model. To better understand how environmental conditions affect tolerance to herbivory, the effects of compensatory growth should be taken into account.     

Nitrogen cycling in poplar stands defoliated by insects

Large-scale outbreaks of defoliating insects are common in temperate forests. These outbreaks are thought to be responsible for substantial cycling of nitrogen (N), and its loss from the system. Gypsy moth (Lymantria dispar) populations within poplar plots were manipulated over 2 years so that the ecosystem-wide consequences of catastrophic defoliation on N cycling could be examined. The quantities of N in leaf litter-fall, ammonia volatilization and soil N pools were estimated across the two seasons. Defoliated leaf biomass was estimated from experimentally derived approximate digestibility factors and added to the mass of senesced leaf to determine total annual leaf production. Throughout the growing season the defoliation treatment peaked at about 40% in year 1 and 100% in year 2. Rapid regrowth after defoliation meant that only 45% of the annual leaf biomass was consumed in the defoliation treatment in year 2, while control plots suffered about 20% consumption each year. In each year, defoliated plots produced 20% more leaf biomass and N than the controls, a phenomenon attributed to compensatory photosynthesis. No substantial losses of N via ammonia volatilization, nitrous oxide emission or nitrate leaching were observed. Neither was there any sustained or substantial gain in the soils microbial biomass or inorganic N pools. These observations suggest that the defoliated poplars were able to compete with soil microbes and N loss mechanisms for soil N as it became available, thereby ameliorating the effects of defoliation on soil nitrogen cycling. We conclude from this study that the N mineralized from defoliation residues was conserved in this plantation ecosystem.     

Seed germination and seedling survival traits of invasive and non-invasive congeneric <Emphasis Type="Italic">Ruellia</Emphasis> species (Acanthaceae) in Yucatan,Mexico

We compared requirements for seed germination and seedling establishment for Ruellia nudiflora, an invasive species in Yucatan, Mexico with those of the congeneric non-invasive R. pereducta. Germination and seedling survival rates were higher for R. nudiflora than for R. pereducta under high light. Additionally, the ranges of temperature and water potential that allow germination for R. nudiflora were much broader than those of R. pereducta. Seedlings of R. nudiflora exhibited higher survival to drought by shedding their leaves during drought, an important strategy in environments under extreme drought. Seedlings of R. nudiflora also exhibited higher extreme temperature tolerance than R. pereducta seedlings. Overall, traits exhibited by R. nudiflora such as ability to germinate under a wide range of conditions, adaptation to environmental stress and high tolerance to environmental heterogeneity during the seedling stage, have been repeatedly recognized as determinants of colonization success of invasive species in open disturbed areas.     

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.     

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.     

Lack of compensatory growth under phosphorus deficiency in grazing-adapted grasses from the Serengeti Plains

Summary Two shortgrass species (Sporobolus ioclados and Eustachys paspaloides) and two midgrass species (E. paspaloides and Pennisetum mezianum) from the Serengeti grasslands of Tanzania were grown under conditions of extreme phosphorus (P) deficiency. Production of each of these species is maintained or enhanced by defoliation under adequate nutrient supply (McNaughton et al. 1983). However, under the P-deficient conditions of our experiment, defoliation caused a reduction in biomass of all plant parts of each species. Green leaf biomass was reduced most strongly by defoliation, and crowns were least affected. Yield of biomass and nutrients to grazers (green leaves+clipped material) was enhanced by weekly defoliation in the shortgrass grazing-adapted species, whereas yield to producers (live biomass and nutrients retained by the plant) and yield to decomposers (litter) were strongly reduced by defoliation in all species. Phosphate absorption capacity (V _max) measured on excised roots was enhanced by defoliation in the grazing-adapted Sporobolus, but, due to low affinity (high K _m) of roots of defoliated plants for phosphate, absorption rate was not greatly altered at low solution concentrations. Phosphate absorption capacity was reduced or unaffected by defoliation in other species. We conclude that under conditions of P deficiency, plants are unable to acquire the nutrients necessary to replenish large nutrient losses to grazers. In low-nutrient environments, compensatory growth (stimulation of production by grazing) is not a viable strategy. Therefore, in these environments plants respond evolutionarily to herbivores by developing chemical or morphological defenses.     

Response of an invasive liana to simulated herbivory: implications for its biological control

Pre-release evaluation of the efficacy of biological control agents is often not possible in the case of many invasive species targeted for biocontrol. In such circumstances simulating herbivory could yield significant insights into plant response to damage, thereby improving the efficiency of agent prioritisation, increasing the chances of regulating the performance of invasive plants through herbivory and minimising potential risks posed by release of multiple herbivores. We adopted this approach to understand the weaknesses herbivores could exploit, to manage the invasive liana, Macfadyena unguis-cati. We simulated herbivory by damaging the leaves, stem, root and tuber of the plant, in isolation and in combination. We also applied these treatments at multiple frequencies. Plant response in terms of biomass allocation showed that at least two severe defoliation treatments were required to diminish this liana's climbing habit and reduce its allocation to belowground tuber reserves. Belowground damage appears to have negligible effect on the plant's biomass production and tuber damage appears to trigger a compensatory response. Plant response to combinations of different types of damage did not differ significantly to that from leaf damage. This suggests that specialist herbivores in the leaf-feeding guild capable of removing over 50% of the leaf tissue may be desirable in the biological control of this invasive species.     

Differential effects of type and quantity of leaf damage on growth,reproduction and defence of lima bean (Phaseolus lunatus L.)

Folivores are major plant antagonists in most terrestrial ecosystems. However, the quantitative effects of leaf area loss on multiple interacting plant traits are still little understood. We sought to contribute to filling this lack of understanding by applying different types of leaf area removal (complete leaflets versus leaflet parts) and degrees of leaf damage (0, 33 and 66%) to lima bean (Phaseolus lunatus) plants. We quantified various growth and fitness parameters including above‐ and belowground biomass as well as the production of reproductive structures (fruits, seeds). In addition, we measured plant cyanogenic potential (HCNp; direct chemical defence) and production of extrafloral nectar (EFN; indirect defence). Leaf damage reduced above‐ and belowground biomass production in general, but neither variation in quantity nor type of damage resulted in different biomass. Similarly, the number of fruits and seeds was significantly reduced in all damaged plants without significant differences between treatment groups. Seed mass, however, was affected by both type and quantity of leaf damage. Leaf area loss had no impact on HCNp, whereas production of EFN decreased with increasing damage. While EFN production was quantitatively affected by leaf area removal, the type of damage had no effect. Our study provides a thorough analysis of the quantitative and qualitative effects of defoliation on multiple productivity‐related and defensive plant traits and shows strong differences in plant response depending on trait. Quantifying such plant responses is vital to our understanding of the impact of herbivory on plant fitness and productivity in natural and agricultural ecosystems.     

Partial compensation in Psychotria marginata (Rubiaceae) after simulated defoliation increases photon capture and photosynthesis

We used Y-plant, a computer-based model of plant crown architecture analysis, to simulate effects of defoliation on daily canopy carbon gain in Psychotria marginata (Rubiaceae) plants under two contrasting irradiances. Five levels of defoliation were simulated using two different types of leaf blade damage. Compensatory increases in photon-saturated photosynthetic capacity (P _max) of 25, 50, and 100 % defoliation were also simulated. In all simulations daily photon capture and CO₂ assimilation increased with defoliation. However, without a compensatory response, daily canopy carbon gain also decreased with defoliation. Under high irradiance, reduction in daily canopy carbon gain was less than what would be expected if the response was proportional to leaf area reduction. Thus, 25 and 50 % defoliation resulted in only 20 and 41 % of daily canopy carbon gain reduction, respectively. In the scenario where 25 % of the leaf area was removed, if the P _max value was increased by 25 %, the remaining leaves compensated for 94 % of the daily canopy carbon relative to an undamaged non-compensated plant. At the same defoliation level, incrementing P _max values by 50 and 100 % resulted in overcompensation. Hence, because the increment of daily photon capture and CO₂ assimilation after defoliation was more a passive consequence of the reduction in leaf area than an active response, under the conditions tested photosynthetic compensation could be only possible through an active mechanism such as the increment of P _max values.     

Effect of stress and time for recovery on the amount of compensatory growth after grazing

Summary We tested the hypothesis that the amount of compensatory growth after defoliation is affected by the level of stress at which plants grow when defoliated and by the length of time for recovery. Growth response to defoliation went from partial compensation when plants were growing at high relative growth rates (RGR) to overcompensation when plants were more stressed and growing at low RGR. Defoliation released plants from the limitation imposed by the accumulation of old and dead tissue and this release overrode the negative effect of biomass loss. Compensatory growth resulted from a higher RGR aboveground that was not associated with a reduction in RGR belowground. Time available for recovery had a major impact on the outcome of defoliation. With a short time for recovery, RGR was decreased by defoliation because an immediate increase in net assimilation rate was overridden by a reduction in the ratio of leaf area to plant weight. After defoliation, this ratio increased quickly due to a larger allocation to leaf growth and lower leaf specific weights, resulting in higher RGR. We conclude that the compensatory response to grazing depends on the type and level of stress limiting growth. Allocation and physiological responses to stress may positively or negatively affect the response to grazing and, simultaneously, grazing may alleviate or aggravate the effects of different types of stress.     

Effects of leaf beetle damage on stem wood production in coppicing willow

1 The effect of defoliation by larvae of the leaf beetle Phratora vulgatissima on current‐year stem wood production of resprouting Salix viminalis was investigated for two years. Adjacent subplots with varying levels of defoliation within one large willow plantation in south Sweden were studied in the two years. 2 High defoliation levels reduced stem wood production by an average of 32 and 39% in the two years, respectively. 3 Medium defoliation levels reduced stem wood production by 16% in one year. In the other year, the stem wood production of medium‐defoliated stools did not differ significantly from stools exposed to low defoliation, i.e. there was full compensatory growth. 4 The main difference between the year with compensatory growth and the one without was that overall productivity was higher in the year with compensation. This finding forms the basis for a mechanistic model by which compensation could be accomplished. We propose that the major contribution to full compensation comes from an increased growth among intermediate‐sized shoots of medium‐defoliated stools relative to the corresponding shoots in stools exposed to low defoliation.     

植物受动物采食后的补偿作用──影响补偿作用的因素

植物受动物采食后的补偿作用影响补偿作用的因素原保忠王静赵松岭（兰州大学干旱农业生态国家重点实验室,７３００００）ＰｌａｎｔＣｏｍｐｅｎｓａｔｉｏｎｆｏｒＡｎｉｍａｌＨｅｒｂｉｖｏｒｙＦａｃｔｏｒｓＡｆｅｃｔｉｎｇＰｌａｎｔＣｏｍｐｅｎｓａｔｉｏｎ．Ｙ...     

Carbohydrate storage and use in an alpine population of the perennial herb, Oxytropis sericea

I tested hypotheses for ecological roles of storage carbohydrates in perennating organs (roots and branches) of alpine Oxytropis sericea, a leguminous herb. In naturally growing plants, total nonstructural carbohydrates achieved their maximal concentration in the fall, declined during winter, and reached minimal levels immediately after growth initiation in the spring. Experimental manipulation of carbon sink-source relations through shading of leaves of reproductive plants revealed that the normally unused portion of these carbohydrates is largely available for withdrawal. In another experiment, plants subjected to carbohydrate depletion through shading suffered decreased leaf growth after winter dormancy and had a lower probability of flowering and decreased inflorescence biomass. The dependence of reproductive growth on stored carbohydrates, however, was limited to its initial stages, because accumulation of storage carbohydrates occurred simultaneously with inflorescence expansion, flowering, and fruiting. Moreover, the whole-plant photosynthetic rate, estimated from gas exchange measurements also peaked at the time of inflorescence growth. To address whether stored reserves allow compensatory regrowth following defoliation, plants were subjected to experimental removal of leaves and inflorescences. Defoliated O. sericea partly regrew the lost leaves but withdrawal of stored carbohydrates was limited. Similarly, in a second defoliation experiment where infructescences were left intact, the plants used little stored carbohydrate and only partly compensated for fruit growth. However, carbohydrate accumulation was negatively affected by defoliation. While the ecological importance of stored nonstructural carbohydrates cannot be attributed to any function in isolation, winter respiration, leaf regrowth after winter, and early reproductive growth in O. sericea all depend to a significant extent on stored reserves. Maintaining a large storage pool may protect these functions in years when carbon status is less favorable than during this study. Received: 13 May 1998 / Accepted: 24 November 1998