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.     

Clonal integration and effects of simulated herbivory in old-field perennials

Summary We compared the growth, phenology and leaf demography of partly defoliated, connected shoots with that of partly defoliated, severed shoots in four old-field perennials (Solidago canadensis, S. altissima, S. gigantea, Aster lanceolatus) with differing genet architectures (rhizome systems), in a common garden and in the field. Our main hypothesis was that defoliation would have fewer negative effects on shoot performance if shoots were connected than if their rhizomes were severed. Since degree of clonal integration is related to differences in genet architecture, our second hypothesis was that the effects of defoliation would be less pronounced in more integrated than in less integrated clones. Removing about 50% of the total leaf area from shoots had different effects depending on plant species, shoot density, and in particular whether rhizome connections between shoots were left intact or severed. In agreement with our prediction, experimentally isolated shoots in the field or in high density clumps in the garden suffered the most from defoliation, while shoots with intact connections or in low density clumps suffered the least. Our second prediction was neither confirmed nor falsified in the present study. Solidago altissima showed overcompensation in response to simulated herbivory in the common garden, i.e. defoliated shoots grew faster and were larger at harvest than their non-defoliated neighbours.     

Carbon autonomy of reproductive shoots of Siberian alder (<Emphasis Type="Italic">Alnus hirsuta</Emphasis> var.<Emphasis Type="Italic"> sibirica</Emphasis>)

Carbon autonomy of current-year shoots in flowering, and of current-year shoots plus 1-year-old shoots (1-year-old shoot system) in fruiting of Siberian alder (Alnus hirsuta var. sibirica) was investigated using a stable isotope of carbon, ¹³C. The current-year shoot and 1-year-old shoot systems were fed ¹³CO₂ and the atom% excess of ¹³C in flowers and fruits was determined. The majority of photosynthate allocated to flower buds was originally assimilated in the leaves of the flowering current-year shoots. Of all the current-year shoots on fruiting 1-year-old shoots, only those nearest to the fruits allocated the assimilated photosynthate to fruit maturation. These results indicate that the current-year shoots and 1-year-old shoot systems are carbon-autonomous units for producing flowers and maturing fruits, respectively.     

Growth,chemical responses and herbivory after simulated leaf browsing in <Emphasis Type="Italic">Combretum apiculatum</Emphasis>

Vegetative and chemical responses to simulated leaf browsing during the growth season, and their subsequent effect on herbivory, were studied on Combretum apiculatum Sonder (Combretaceae) in Botswana. Treatments (50% and 100% leaf and shoot apex removal) were performed just before the shoot growth curve levelled out, and responses recorded 3 months later, just before leaf fall. Compared to controls, defoliation treatments, removing apical dominance, reduced growth in tree height and increased shoot mortality, although the production of lateral shoots increased. At the end of the trial, there was no difference in total length of annual shoots between treatment groups. Significant refoliation occurred only after 100% defoliation. Refoliated leaves were smaller and the 100% defoliated trees had a lower final leaf biomass. Total leaf biomass production was, however, equal for all treatment groups. Refoliated leaves contained higher levels of N, lower levels of acid-detergent fibre (ADF) and total phenolics, and showed a trend towards lower levels of condensed tannins, compared to leaves on control trees. Such chemical changes may be due to either carbon stress or to younger physiological age of new leaves. In spite of the observed potential increase in food quality, we found no evidence of increased levels of insect or ungulate herbivory on refoliated leaves, which, at least for insect herbivory, may be explained by the reduction in temporal availability of leaves. We conclude that the single severe defoliation was not detrimental to C. apiculatum in the short-term, although the resource loss and induced compensatory growth may produce negative effects during subsequent growth seasons.     

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.     

Leaf age and the timing of leaf abscission in two tropical dry forest trees

We used experimental defoliations to examine the effect of leaf age on the timing of leaf shedding in two tropical dry forest trees. Trees of the deciduous Bombacopsis quinata (bombacaceae, a.k.a. Pachira quinata) and the brevi-deciduous Astronium graveolens (anacardiaceae) were manually defoliated for three times during the rainy season. All trees started to produce a new crown of leaves 2 weeks after defoliation, and continued expanding leaves throughout the rainy season. At the transition to the dry season, the experimental groups consisted of trees with known differences in maximum leaf age. Defoliations resulted in declines in stem growth but did not affect the mineral content or water relations of the leaves subsequently produced. There was no effect of leaf age on the timing of leaf abscission in B. quinata. In A. graveolens, the initiation of leaf shedding followed in rank order, the maximum leaf age of the four treatments, but there was substantial coherence among treatments in the major period of leaf abscission such that trees completed leaf shedding at the same time. In the two species, leaf water potential (Ψ_L) and stomatal conducantce (g _S) declined with the onset of the dry season, reaching minimum values of –0.9 MPa in P. quinata and <–2.0 MPa in A. graveolens. Within each species, leaves of different age exhibited similar Ψ_L and g _S at the onset of drought, and then decreased at a similar rate as the dry season progressed. Overall, our study suggests that the environmental factors were more important than leaf age in controlling the timing of leaf shedding.     

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.     

Impact of defoliation intensities on plant biomass,nutrient uptake and arbuscular mycorrhizal symbiosis in Lotus tenuis growing in a saline‐sodic soil

The impact of different defoliation intensities on the ability of Lotus tenuis plants to regrowth, mobilise nutrients and to associate with native AM fungi and Rhizobium in a saline‐sodic soil was investigated. After 70 days, plants were subjected to 0, 25, 50, 75 and 100% defoliation and shoot regrowth was assessed at the end of subsequent 35 days. Compared to non‐defoliated plants, low or moderate defoliation up to 75% did not affect shoot regrowth. However, 100% treatment affected shoot regrowth and the clipped plants were not able to compensate the growth attained by non‐defoliated plants. Root growth was more affected by defoliation than shoot growth. P and N concentrations in shoots and roots increased with increasing defoliation while Na⁺ concentration in shoots of non‐defoliated and moderately defoliated plants was similar. Non‐defoliated and moderately defoliated plants prevented increases of Na⁺ concentration in shoots through both reducing Na⁺ uptake and Na⁺ transport to shoots by accumulating Na⁺ in roots. At high defoliation, the salinity tolerance mechanism is altered and Na⁺ concentration in shoots was higher than in roots. Reduction in the photosynthetic capacity induced by defoliation neither changed the root length colonised by AM fungi nor arbuscular colonisation but decreased the vesicular colonisation. Spore density did not change, but hyphal density and Rhizobium nodules increased with defoliation. The strategy of the AM symbiont consists in investing most of the C resources to preferentially retain arbuscular colonisation as well as inoculum density in the soil.     

Carbon allocation in shoots of alpine treeline conifers in a CO<Subscript>2</Subscript> enriched environment

With a new approach we assessed the relative contribution of stored and current carbon compounds to new shoot growth in alpine treeline conifers. Within a free air CO₂ enrichment experiment at the alpine treeline in Switzerland, ¹³C-depleted fossil CO₂ was used to trace new carbon in the two tree species Larix decidua L. and Pinus uncinata Ramond over two subsequent years. The deciduous L. decidua was found to supply new shoot growth (structural woody part) by 46% from storage. Surprisingly, the evergreen P. uncinata, assumed to use current-year photosynthates, also utilized a considerable fraction of storage (42%) for new wood growth. In contrast, the needles of P. uncinata were built up almost completely from current-year photosynthates. The isotopic composition of different wood carbon fractions revealed a similar relative allocation of current and stored assimilates to various carbon fractions. Elevated CO₂ influenced the composition of woody tissue in a species-specific way, e.g. the water soluble fraction decreased in pine in 2001 but increased in larch in 2002 compared to ambient CO₂. Heavy defoliation applied as an additional treatment factor in the second year of the experiment decreased the lipophilic fraction in current-year wood in both species compared to undefoliated trees. We conclude that storage may play an important role for new shoot growth in these treeline conifers and that altered carbon availability (elevated CO₂, defoliation) results in significant changes in the relative amount of mobile carbon fractions in woody tissue. In particular, stored carbon seems to be of greater importance in the evergreen P. uncinata than has been previously thought.     

Development of shoot inHydrangea macrophylla II. sequence and timing

The development of axillary buds, terminal buds, and the shoots extended from them was studied inHydrangea macrophylla. The upper and lower parts in a nonflower-bearing shoot are discernible; the preformed part of a shoot develops into the lower part and the neoformed part into the upper part (Zhou and Hare, 1988). These two part are formed by the different degrees of internode elongation at early and late phases during a growth season, respectively. Leaf pairs in the neoformed part of the shoot are initiated successively with a plastochron of 5–20 days after the bud burst in spring. The upper axillary buds are initiated at approximately the same intervals as those of leaf pairs, but 10–30 days later than their subtending leaves. Changes in numbers of leaf pairs and in lengths of successive axillary buds show a pattern similar to the changes in internode lengths of the shoot at the mature stage. The uppermost axillary buds of the flower-bearing shoot often begin extending into new lateral shoots when the flowering phase has ended. The secondary buds in terminal and lower axillary buds are initiated and developed in succession during the late phase of the growth season. Internode elongation seems to be important in determining the degrees of development of the axillary buds. Pattern of shoot elongation is suggested to be relatively primitive. Significances of apical dominance and environmental conditions to shoot development are discussed.     

Indirect shoot organogenesis from leaves of Dieffenbachia cv. Camouflage

A novel protocol for indirect shoot organogenesis of Dieffenbachia cv. Camouflage was established using leaf explants excised from in vitro shoot cultures. The frequency of callus formation reached 96% for explants cultured on Murashige and Skoog (1962) basal medium supplemented with 5 μM thidiazuron and 1 μM 2,4-dichlorophenozyacetic acid. The number of shoots regenerated was high, with up to 7.9 shoots produced per callus cultured on basal medium supplemented with 40 μM N ⁶-(Δ²-isopentenyl)adenine and 2 μM indole-3-acetic acid. Regenerated shoots rooted well in a soilless substrate, acclimatized ex vitro at 100%, and grew vigorously under shaded greenhouse conditions. Somaclonal variations in leaf variegation, color, and morphology have been observed in regenerated plants.     

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

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

Early spring defoliation,secondary leaf flush,and leafminer outbreaks on American holly

Summary Adults of Phytomyza ilicicola (Diptera: Agromyzidae), a univoltine specialist leafminer, emerge in close synchrony with leaf flush of American holly and feed on and oviposit in soft, partially expanded leaves. Early spring defoliation, such as commonly results from freezing injury to young shoots, is followed several weeks later by a second flush of young leaves from lateral buds. We simulated this phenomenon by manually defoliating whole small trees and individual shoots of large trees to test the hypothesis that freezing injury can encourage leafminer outbreaks by inducing an abundance of soft, protein rich young leaves late in the adult activity period, when availability of vulnerable leaves becomes limited. Defoliation of small trees one or two weeks after bud break resulted in six- to 13-fold increases in the incidence of feeding punctures and larval mines on second flush leaves as compared with densities on original young leaves of control trees. Similarly, we induced significant increases in feeding punctures and larval mines on second flush leaves of individual defoliated shoots, although leaves that did not open until after the flight period escaped this injury. These observations underscore the capability of adult female P. ilicicola to locate and exploit a small number of phenologically available leaves among many hundreds of older leaves on the same tree. By altering the phenology of leaf flush, certain kinds of environmental stress may predispose perennial plants to outbreaks of early season folivores that restrict their feeding or oviposition to very young leaves.     

Current-year shoot based approach for annual variation in the reproductive output in Siberian alder (<Emphasis Type="Italic">Alnus hirsuta</Emphasis> var. <Emphasis Type="Italic">sibirica</Emphasis>)

Siberian alder (Alnus hirsuta var. sibirica) shows annual variation in reproductive output. However, this phenomenon has not been explained by hypotheses proposed in previous studies. In this study, we constructed a matrix model of current-year shoot dynamics to estimate and compare the reproductive output of current-year shoot population with or without annual variation, and explained the annual variation of reproductive output in terms of a reproductive strategy of the current-year shoot population of Siberian alder. The current-year shoot population of Siberian alder was divided into three functional groups: reproductive, maintenance, and explorative shoot sub-populations. A transition matrix was calculated from the relationships between 1-year-old shoots and the current-year shoots on them. The dynamics of the current-year shoot population was simulated using the estimated matrix on the patterns with or without annual variation in reproductive output. The pattern with annual variation in reproductive output yielded more reproductive current-year shoots than the pattern with high reproductive output every year. The annual variation of reproductive output may well be regarded as a reproductive strategy of current-year shoots to increase lifetime fecundity.     

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

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

An improved micropropagation of <Emphasis Type="Italic">Eclipta alba</Emphasis> by in vitro priming with chlorocholine chloride

An efficient method of micropropagation for Eclipta alba from young nodal axils of shoot tip explants has been developed by giving special attention to ‘priming’ in vitro plantlets in view of increasing their hardening ability after transplantation ex vitro. Among 3 cytokinins—BAP, kinetin and TDZ, BAP was found most effective in inducing and proliferating adventitious shoots. The highest frequency of responding explants (100%) and maximum number of shoots (23.0) per explant were obtained after 60 days culture on MS medium containing 8.8 μM BAP. Cent percent shoots developed roots directly from shoot base when transferred to growth regulator-free MS medium. For priming E. alba microshoots, 6.3 μM of chlorocholine chloride (CCC) was found most effective. The major changes observed in 30 days old treated shoots were, production of increased number of root, elevation of chlorophyll level in leaves and increase in plant biomass. Furthermore, arrested undesirable shoot elongation made the plants sturdier and more suitable for acclimatization. The primed micropropagated E. alba plants were healthy and survived by higher frequency (100%) in soil in comparison to the non-treated plants (84% survival).     

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.     

Morphogenesis induction and organogenic nodule differentiation in <Emphasis Type="Italic">Populus euphratica</Emphasis> Oliv. leaf explants

Populus euphratica Oliv. is a deciduous poplar species, occurring mainly in riparian areas of China and Middle Eastern countries, and presenting high tolerance to extreme temperatures and high soil salinity. In this study, an optimized protocol for development and propagation of P. euphratica from leaf explants is reported, based on a morphogenic process that involves organogenic nodule differentiation. Adventitious shoot regeneration of P. euphratica from organogenic nodules of leaf explants was achieved within a range of concentrations of α-naphtalenacetic acid and 6-benzylaminopurine, at a fixed 2:1 ratio. Cambial cells started to divide 5 days after inoculation on culture medium and, after 12 days, several organizing centres were already formed. Non-friable callus tissue, together with organization centres, formed structures that evolved to nodules after about 40 days which were, then, able to regenerate new shoots after 50–60 days. The nodules did not separate from the mother explants and were able to successfully give rise to new adventitious shoots. These were rescued and successfully grown and rooted in different culture media, and fully developed plants were obtained. The regeneration system here described for P. euphratica is innovative, reproducible and data from histological studies of the morphogenic process support the classification of the regenerative structures as organogenic nodules.     

Endophytic and epiphytic phyllosphere fungi of red-osier dogwood (<Emphasis Type="Italic">Cornus stolonifera</Emphasis>) in British Columbia

Endophytic and epiphytic phyllosphere fungi associated with red-osier dogwood (Cornus stolonifera), a deciduous shrub, were examined in coastal British Columbia, Canada. Current-year shoots were divided into four types based on the absence or presence of inflorescence and secondary elongated shoots at the apex of primary shoots. Leaves on these shoots were then classified into six categories so as to examine the effect of flowering, secondary shoot elongation, and shoot order within current-year shoots on the occurrence of phyllosphere fungi. Species composition of fungi was markedly different between the interior and surface of leaves, whereas it was relatively similar among the six leaf categories in the interior or on the surface. Frequencies of the eight major species were not different between leaves on flowering and nonflowering shoots. The frequency of Colletotrichum gloeosporioides in the leaf interior was greater on leaves on the primary shoots that elongated the secondary shoots than on those that did not, and was greater on leaves on the primary shoots than on those on the secondary shoots. On the other hand, secondary shoot elongation and shoot order had no effect on the frequencies of C. gloeosporioides and the other seven epiphytes on leaf surfaces.     

Crown hollowing as a consequence of early shedding of leaves and shoots

Leafing pattern has long been considered as an important element characterizing the growth strategy of tree species; however, the consequences of leafing pattern for tree-crown formation have not been fully understood. To address this issue, the dynamic events (growth, birth, and death) of current-year shoots and leaves were investigated together with their location in saplings of a pioneer tree, Alnus sieboldiana. The leafing pattern was characterized by successive emergence and shedding of short-lived leaves. The combination of successive leafing and within-crown variation in leaf production brought about characteristic outcomes in crown morphology. In the outer crown, because of continuous leaf production, the shoots achieved great extension and enormous daughter shoot production, resulting in rapid expansion of the crown. In contrast, in the inner crown, due to early termination of leaf production, the shoots completely lost their leaves early in the growing season and consequently themselves died and were shed within the season. Such quick shedding of shoots caused “crown hollowing”, i.e., the interior crown consisted of primary branches with little secondary development or foliage. These dynamic features are an effective adaptive strategy in early succession but also may be a disadvantage to maintaining foliage for longer period. Crown maintenance associated with the longevity of structural components is thought to play an important role in survival strategy of tree species.