Growth Rate and Water Relations of Citrus Leaf Flushes

Growth rates of seasonal leaf flushes of ‘Valencia’orange [Citrus sinensis (L.) Osbeck] were measured and waterrelations characteristics of young (new) and over-wintered (old)citrus leaves were compared. New flush leaves had lower specificleaf weights and lower midday leaf water potentials than comparablyexposed old leaves. Spring and summer flush new leaves had higherosmotic potentials than old leaves. These differences becamenon-significant as the new leaves matured. During summer conditions,water-stressed new leaves reached zero turgor and stomatal conductancealso began to decrease in them at higher leaf water potentialsthan in old leaves. Old leaves were capable of maintaining openstomata at lower leaf water potentials. Opened flowers and newflush leaves lost more water, on a dry weight basis, than flowerbuds, fruit or mature leaves. The results illustrate differencesin leaf water potential and stomatal conductance which can beattributed to the maintenance of leaf turgor by decreases inleaf osmotic potentials as leaves mature. These changes in citrusleaf water relations are especially important since water stressresulting from high water loss rates of new tissues could reduceflowering and fruit set. Citrus sinensis (L.) Osbeck, orange, Citrus paradisi Macf., grapefruit, growth rate, leaf water relations, osmotic potential, water potential, stomatal conductance     

Citrus Leprosis and its Status in Florida and Texas: Past and Present

According to published reports from 1906 to 1968, leprosis nearly destroyed the Florida citrus industry prior to 1925. This was supported with photographs showing typical leprosis symptoms on citrus leaves, fruit, and twigs. Support for the past occurrence of citrus leprosis in Florida includes: (1) presence of twig lesions in affected orange blocks in addition to lesions on fruits and leaves and corresponding absence of similar lesions on grapefruit; (2) yield reduction and die-back on infected trees; and (3) spread of the disease between 1906 and 1925. Transmission electron microscopy (TEM) examination of tissue samples from leprosis-like injuries to orange and grapefruit leaves from Florida in 1997, and fruits from grapefruit and sweet orange varieties from Texas in 1999 and 2000 did not contain leprosis-like viral particles or viroplasm inclusions. In contrast, leprosis viroplasm inclusions were readily identified by TEM within green non-senescent tissues surrounding leprosis lesions in two of every three orange leaf samples and half of the fruit samples obtained from Piracicaba, Brazil. Symptoms of leprosis were not seen in any of the 24,555 orange trees examined across Florida during 2001 and 2002. The authors conclude that citrus leprosis no longer exists in Florida nor occurs in Texas citrus based on: (1) lack of leprosis symptoms on leaves, fruit, and twigs of sweet orange citrus varieties surveyed in Florida: (2) failure to find virus particles or viroplasm inclusion bodies in suspect samples from both Florida and Texas examined by TEM; (3) absence of documented reports by others on the presence of characteristic leprosis symptoms in Florida; (4) lack of its documented occurrence in dooryard trees or abandoned or minimal pesticide citrus orchard sites in Florida. In view of the serious threat to citrus in the U.S., every effort must be taken to quarantine the importation of both citrus and woody ornamental plants that serve as hosts for Brevipalpus phoenicis (Geijskes), B. californicus (Banks), and B. obovatus Donnadieu (Acari: Tenuipalpidae) from countries where citrus leprosis occurs.     

Novel expression patterns of carotenoid pathway-related genes in citrus leaves and maturing fruits

Carotenoids are abundant in citrus fruits and vary among cultivars and species. In the present study, high performance liquid chromatography (HPLC) and real-time polymerase chain reaction (PCR) were used to investigate the expression patterns of 23 carotenoid biosynthesis gene family members and their possible relation with carotenoid accumulation in fresh flavedo, juice sacs and leaves of Valencia orange during fruit maturation. Violaxanthin and lutein mainly accumulated in fruit (flavedo and juice sacs) and leaves (young and mature), respectively, accounting for nearly 79 %, 57 %, 53 % and 70 % of corresponding total carotenoids in February. Violaxanthin content quickly began to increase in flavedo in December, but the increase in juice sacs began later in January. In mature leaves, lutein content was three times that in young leaves; α-carotene and β-carotene were also much higher in mature leaves than in flavedo or juice sacs. Generally most of the carotenoid biosynthesis gene members were expressed at higher levels in flavedo than in juice sacs, and the expression of some continued to increase in flavedo during fruit maturation. All CHYB members expressed at high levels and had similar patterns in juice sacs. Interestingly, the capsanthin capsorubin synthase (CCS) members had similar expression levels and patterns in flavedo and juice sacs. Differences in gene expression between leaf and fruit tissues were noted, pointing to some tissue specificity for certain members of the gene families associated with carotenogenesis. The expression patterns of these 23 citrus carotenoid biosynthesis gene members were also compared with their expression patterns in other plants. Taken together, these first-hand expression data will be useful to define the tissue-specific roles of each gene member in accumulation of different carotenoids in citrus leaves and maturing fruits.     

Comparative response to citrus foliage and citrus fruit odour by wild and mass-reared sterile Mediterranean fruit fly males of a genetic sexing strain

In greenhouse behavioural assessments conducted at Seibersdorf, Austria that allowed flies the choice of different types of potted host trees and plants, it was found that mature laboratory mass-reared Mediterranean fruit fly males, Ceratitis capitata (Wied), (Dipt. Tephritidae), strain Vienna-42 (temperature-sensitive lethal genetic sexing strain) showed the same preference for orange foliage, over other plants types and parts, as wild males. They aggregated in leks, released pheromone, and courted approaching females almost exclusively on orange foliage. Even though a number of other trees and plants were available to Vienna-42 males, less than 1% of these sexual activities occurred on the foliage or other plant parts of non-Citrus trees and not a single mating occurred there. In choice tests carried out in Chios, Greece in large field cages housing naturally planted orange trees, it was determined that mature and sterile Mediterranean fruit fly males of the same sexing strain responded to the odour emanating from the flavedo (due to superficial cuts made in the fruit peel containing the oil glands) of ripening sweet, sour oranges and lemons, in a similar intensity, order of fruit preference and time of day as did mature males of the local wild population. Males of both strains were frequently observed contacting with their mouth parts the oils exuding from the cuts made in the flavedo of the orange peel, or exhibiting homosexual activity on, or near the cuts. The results show that mass-reared, sterile males of the Vienna-42, genetic sexing strain behave in a comparable way to wild males in an important component of habitat location (odour of host foliage and host fruit) and hence are competitive with them in these respects. None of the standard international quality control tests for sterile Mediterranean fruit flies evaluates this important habitat location behaviour, which is a prerequisite for the successful encounter of the sexes in a sterile release programme.     

Population dynamics ofHemisarcoptes coccophagus Meyer (Astigmata: Hemisarcoptidae) attacking three species of armored scale insects (Homoptera: Diaspididae)

The parasitic miteHemisarcoptes coccophagus Meyer (Astigmata: Hemisarcoptidae) feeds on armored scale insects (Homoptera: Diaspididae). Three host scales were studied: chaff scale (Parlatoria pergandii Comstock andP. cinerea Hadden) infesting citrus (grapefruit and orange) orchards, latania scale [Hemiberlesia lataniae (Signoret)] and olcander scale (Aspidiotus nerii Bouche) occurring on acacia (Acacia cyanophylla) branches and leaves. The overall dynamics of mite populations were similar on the different scale species, although the level of attack on each sampled diaspidid host was different. Rates of young mite stages were highly variable on all three hosts throughout the sampling period while adult rates were stable, constituting around 20% of all mites. No oviposition by mites occurred in mid-winter on any scale species, on any plants or parts, at all sampling sites. Of the three diaspidids, highest mite prevalence (parasitization rate) occurred on latania scale. Mite prevalence on oleander scale infesting mature leaves was higher than on young leaves.H. coccophagus apparently first attacks latania scale on branches, it then disperses to the same host inhabiting mature leaves, from there to co-occurring oleander scale and finally to oleander scale infesting young leaves. Mite prevalence on chaff scale was influenced by host plants and climatic regions: in the Negev it was higher than in the coastal plain, with scales on grapefruit being more heavily attacked at the former site but not on coastal plain. Our data suggest thatH. coccophagus prefers ovipositing scale females. The decreasing order of preference for other scale stages was: young females immature males, 2nd-instar nymphs females.     

The effects of plant quality on caterpillar growth and defense against natural enemies

A survey of 85 species of Lepidoptera feeding on 40 hosts on Barro Colorado Island, Panama showed that growth and defensive traits of caterpillars were correlated with the nutritional and defensive traits of their hosts. Growth rates were faster on young than mature leaves, reflecting the higher nitrogen and water content of the former. Growth was also positively correlated with leaf expansion rate, partially because of higher nitrogen and water contents of fast-expanding young leaves. Specialists grew faster than generalists, but both responded positively to nutritional quality. There was no effect of lepidopteran family on growth. In analyses where the effects of nitrogen and water were removed, the residuals for growth rate were greater for young than for mature leaves and were positively correlated with expansion rates of young leaves. This suggests that traits other than nutrition were also important. As young, expanding leaves cannot use toughness as a defense, one possible explanation for the differences in growth is differences in chemical defenses. Growth rate residuals for both specialists and generalists were higher for the more poorly defended fast-expanders, but the effect was greatest for generalists, perhaps because generalists were more sensitive to secondary metabolites. We predicted that slow growth for caterpillars would increase their risk to natural enemies and would select for higher defenses. Generalists had more defensive traits than specialists and were less preferred in feeding trials with ants. Similarly, species feeding on mature leaves were the most defended and those feeding on fast-expanding young leaves were the least defended and most preferred by ants. Thus the effects of plant secondary metabolites and nutrients dictate herbivore growth rates, which in turn influence their susceptibility to the third trophic level and the importance of defenses.     

Demographic consequences of chromatic leaf defence in tropical tree communities: do red young leaves increase growth and survival?

Background

Many tropical forest tree species delay greening their leaves until full expansion. This strategy is thought to provide newly flushing leaves with protection against damage by herbivores by keeping young leaves devoid of nutritive value. Because young leaves suffer the greatest predation from invertebrate herbivores, delayed greening could prevent costly tissue loss. Many species that delay greening also produce anthocyanin pigments in their new leaves, giving them a reddish tint. These anthocyanins may be fungicidal, protect leaves against UV damage or make leaves cryptic to herbivores blind to the red part of the spectrum.

Methods

A comprehensive survey was undertaken of seedlings, saplings and mature trees in two diverse tropical forests: a rain forest in western Amazonia (Yasuní National Park, Ecuador) and a deciduous forest in Central America (Barro Colorado Island, Panamá). A test was made of whether individuals and species with delayed greening or red-coloured young leaves showed lower mortality or higher relative growth rates than species that did not.

Key results

At both Yasuní and Barro Colorado Island, species with delayed greening or red young leaves comprised significant proportions of the seedling and tree communities. At both sites, significantly lower mortality was found in seedlings and trees with delayed greening and red-coloured young leaves. While there was little effect of leaf colour on the production of new leaves of seedlings, diameter relative growth rates of small trees were lower in species with delayed greening and red-coloured young leaves than in species with regular green leaves, and this effect remained when the trade-off between mortality and growth was accounted for.

Conclusions

Herbivores exert strong selection pressure on seedlings for the expression of defence traits. A delayed greening or red-coloured young leaf strategy in seedlings appears to be associated with higher survival for a given growth rate, and may thus influence the species composition of later life stages.     

Polyamine biosynthesis and titer during various developmental stages of Phaseolus vulgaris

The activities of arginine decarboxylase (ADC; EC 4.1.1.19) and ornithine decarboxylase (ODC; EC 4.1.1.17) as well as polyamine content were examined in Phaseolus vulgaris L. cv. Taylor's Horticultural before and during anthesis, during fruit development and throughout vegetative growth. The specific activities of polyamine biosynthetic enzymes were highest in all rapidly growing tissues, e.g., root apices, hypocotyls, young internodes, young leaves, flower buds, young pods and pericarps. They were lowest in mature, non-growing tissues. Similarly, the content of the major polyamines (putrescine, spermidine, spermine) is highest in rapidly growing tissues, and lowest in mature tissue. These correlations reinforce the growing connection between polyamines and rates of cell division and metabolic activity during both vegetative and reproductive development.     

Mechanical wounding and abscission in citrus

Fruit detachment force (FDF), ethylene evolution, fruit and leaf drop were determined in Citrus sinensis for periods up to 96 h after mechanical wounding. Injury by removing a thin section of mature fruit flavedo reduced FDF, increased ethylene evolution and promoted abscission. Injuring flavedo 1 cm below the calyx was more effective at reducing FDF than injuring flavedo at the equator or the blossom‐end of mature fruit. Injuring the calyx or peduncle of mature fruit, or injuring three leaves closest to the mature fruit did not reduce FDF. Immature fruitlets either did not abscise or underwent low rates of abscission in response to mechanical wounding, depending on age. Inhibiting ethylene binding in wounded mature fruit with 1‐methylcyclopropene (1‐MCP) increased ethylene evolution compared with wounded fruit alone, but the reduction in FDF was similar. When an ethylene biosynthesis inhibitor (aminoethoxyvinylglycine, AVG) was used, reduction in FDF of wounded mature fruit exposed to AVG was similar to that of wounded fruit alone but ethylene production was markedly reduced. Wounding mature leaf blades in the presence or absence of 1‐MCP resulted in elevated but equal ethylene evolution up to 48 h after wounding, however, no leaf drop occurred. Thereafter, ethylene evolution was higher in 1‐MCP‐treated wounded leaves. Removing up to 77% of the total mature leaf area did not cause leaf drop, nor did wounding tissue across the laminar or petiolar abscission zones. Leaflets of 5 mm length reached nearly 100% abscission after mechanical wounding, whereas wounding leaves 20 mm length resulted in 15% abscission. The data suggest that mechanical wounding of flavedo results in mature fruit abscission, and ethylene binding may not be mandatory to initiate abscission in citrus fruit. The differential response of fruit and leaves at different ages to wounding may be related to potential contribution to carbohydrate accumulation, and production and sensitivity of tissues to an abscission signal(s).     

Brevipalpus californicus, B. obovatus, B. phoenicis, and B. lewisi (Acari: Tenuipalpidae): a Review of their Biology, Feeding Injury and Economic Importance

The genus Brevipalpus includes most of the economically important species of Tenuipalpidae. Many Brevipalpus species reproduce by theletokous parthenogenesis while other species reproduce by male fertilization of female eggs. Previous researchers have determined that Brevipalpus californicus (Banks), B. obovatus Donnadieu, and B. phoenicis (Geijskes) females were haploid with two chromosomes. The life cycle and developmental times for these three species are reviewed. Longevity of each Brevipalpus species is two to three times greater than corresponding longevities of various tetranychid mites. Brevipalpus mites inject toxic saliva into fruits, leaves, stems, twigs, and bud tissues of numerous plants including citrus. Feeding injury symptoms on selected plants include: chlorosis, blistering, bronzing, or necrotic areas on leaves by one or more Brevipalpus mites. Premature leaf drop occurred on 'Robinson' tangerine leaves in Florida (USA). Leaf drop was observed in several sweet orange and grapefruit orchards in Texas (USA) that were heavily infested with Brevipalpus mites feeding on the twigs, leaves, and fruit. Initial circular chlorotic areas appear on both sweet orange and grapefruit varieties in association with developing populations of Brevipalpus mites in Texas. These feeding sites become progressively necrotic, darker in color, and eventually develop into irregular scab-like lesions on affected fruit. Russeting and cracking of the fruits of other plant hosts are reported. Stunting of leaves and the development of Brevipalpus galls on terminal buds were recorded on sour orange, Citrus aurantium L., seedlings heavily infested with B. californicus in an insectary. The most significant threat posed by these mites is as vectors of a potentially invasive viral disease called citrus leprosis.     

Effect of local irradiance on CO(2) transfer conductance of mesophyll in walnut

The acclimation responses of walnut leaf photosynthesis to the irradiance microclimate were investigated by characterizing the photosynthetic properties of the leaves sampled on young trees (Juglans nigraxregia) grown in simulated sun and shade environments, and within a mature walnut tree crown (Juglans regia) in the field. In the young trees, the CO(2) compensation point in the absence of mitochondrial respiration (Gamma*), which probes the CO(2) versus O(2) specificity of Rubisco, was not significantly different in sun and shade leaves. The maximal net assimilation rates and stomatal and mesophyll conductances to CO(2) transfer were markedly lower in shade than in sun leaves. Dark respiration rates were also lower in shade leaves. However, the percentage inhibition of respiration by light during photosynthesis was similar in both sun and shade leaves. The extent of the changes in photosynthetic capacity and mesophyll conductance between sun and shade leaves under simulated conditions was similar to that observed between sun and shade leaves collected within the mature tree crown. Moreover, mesophyll conductance was strongly correlated with maximal net assimilation and the relationships were not significantly different between the two experiments, despite marked differences in leaf anatomy. These results suggest that photosynthetic capacity is a valuable parameter for modelling within-canopies variations of mesophyll conductance due to leaf acclimation to light.     

Evaluation ofMicrothrix inconspicuella, [Lepidoptera: Pyralidae], a potential biological control agent forEmex australis in Australia,carried out in apple orchards in South Africa

Host specificity tests carried out in the laboratory in Australia during 1977, showed thatMicrothrix inconspicuella Ragonot could develop on young apple leaves (Harley et al., 1979). Field studies in unsprayed apple orchards in South Africa showed that some feeding occurred, but fewer than 40% of late instar larvae developed to adults when confined in sleeves on apple tree branches. No feeding or survival occurred in large field cages or in the open. Adults which developed from apple fed larvae were smaller, deformed, occasionally mated and laid fertile eggs but their progeny did not feed or develop on apple fruit or leaves. In conclusion,M. inconspicuella larvae did not develop on apple fruit or leaves in the field, damage was mainly limited to apples already injured and feeding on leaves was minimal. Under normal pest control practicesM. inconspicuella populations did not survive on any part of the apple tree or onE. australis growing under the trees.      

A comparison of the volatile oils of mature versus young leaves of Juniperus scopulorum: Chemosystematic significance

The volatile oils of leaves of clonal, greenhouse grown trees of J. scopulorum cv. platinum were examined from young (juvenile or whip leaves) and mature (adult or scale leaves) foliage. Of 36 compounds subjected to analysis of variance, 19 showed significant differences between young and old leaves. In general, the hydrocarbon terpenoids were higher in concentration in new leaves while the oxygenated compounds were higher in the older leaves. This same pattern has been reported in several other gymnosperms. Numerical taxonomy using data on the volatile oils from young and mature leaves yielded the same relative similarities among the five trees examined. Surprisingly, the plants were more similar in the volatile oil from the young leaves than from the mature leaves. The mixing of different ages of leaves in samples for chemosystematic studies is, however, to be discouraged     

Endo‐1,4‐β‐glucanase gene expression and cell wall hydrolase activities during abscission in Valencia orange

The physiological and molecular events of ethylene‐induced abscission in mature fruit calyx, laminar and floral abscission zones of cv. Valencia orange were examined. Continuous exposure of fruit explants to 5 µl 1⁻¹ ethylene for 2 to 40 h resulted in marked increases in endo‐1,4‐β‐glucanase (cellulase) and polygalacturonase (PG) activities in calyx abscission zones. Two abscission‐related cellulases and one PG were found. The major peak of cellulase activity corresponded to a pI of 8.0 and molecular weight of 51 kDa, whereas the minor cellulase peak had a pI of 5.5. The abscission polygalacturonase had a pI of 5.5. Calyx abscission zone RNA was amplified with degenerate primers based on sequence of the purified Valencia orange calyx abscission cellulase, and cloned. The two partial cellulase cDNA clones were 59% identical at the nucleotide level. Genomic Southern analysis suggested that Valencia orange contained two groups of cellulase genes. A full‐length cDNA clone from each group was isolated from a cDNA library prepared from ethylene‐induced calyx abscission zone mRNA. Both genes were expressed in ethylene‐induced calyx, laminar and floral abscission zones, but were not expressed in non‐induced abscission zones or mature leaves treated with or without ethylene, young bark or young fruit of Valencia.     

Responses of Tylenchulus semipenetrans to Citrus Fruit Removal: Implications for Carbohydrate Competition

Sixteen mature Valencia orange trees on rough lemon rootstock were selected on the basis of approximately equal, naturally occurring populations of Tylenchulus semipenetrans in soil. In March, fruit 1 cm in diameter or less were removed from eight of the trees, which were kept free of fruit for 15 months. In July, 4 months after fruit removal, fibrous root (<2 mm d) mass density of defruited trees was 51% greater and insoluble starch in fibrous roots was 24% less than on control trees with fruit. Female T. semipenetrans per gram of root were 64% more numerous on roots of control trees than on defruited trees at this time. Numbers of female nematodes per tree and of juveniles and males in soil did not differ between treatments 4 months after fruit removal. Root mass density remained higher on defruited than control trees for the remaining 13 months that the trees were studied, while nematode density in soil beneath defruited trees rapidly increased to levels proportionate to the additional root mass density. Nine months after fruit removal (December), starch concentration was 84% higher in roots of defruited trees compared to controls and remained 28% higher than in controls 15 months (May) following fruit removal. Between months 9 and 15 following fruit removal, nematode density in soil beneath defruited trees increased at a rate five times that of nematode density beneath control trees. In May, female fecundity (eggs/female) on defruited trees was 41% greater than on control trees. The data were consistent with the hypothesis that carbohydrate competition between developing citrus fruit and T. semipenetrans influences seasonal fluctuations in nematode population densities.     

INFLORESCENCE TYPES AND FRUITING PATTERNS IN HAMLIN AND VALENCIA ORANGES AND MARSH GRAPEFRUIT

The structural, flowering, and fruit-setting patterns of inflorescences of mature Hamlin and Valencia orange and Marsh grapefruit trees were studied for three years. Several development patterns were found, some of which were relatively consistent for the different varieties or years of study. The sequence of anthesis on an inflorescence was: apical flower first, then basal flower, then the subapical flower. Terminals on which the earlier flowers appeared tended to have more inflorescences than those on which flowers appeared later. Inflorescences on which earlier flowers appeared also produced more flowers than those which began flowering later. Inflorescences that began flowering later were more likely to have leaves or have a greater number of leaves than earlier inflorescences. More than half of the inflorescences carried no leaves, and most of these had one flower. No evidence of a relationship between number of flowers and length of the inflorescence was found. Fruit set occurred primarily during the latter part of the flowering period. Many fruit were set on inflorescences without leaves, but on the basis of percent of flowers setting fruit, inflorescences with leaves were more productive. The greatest fruit set occurred in the subapical position on the inflorescence. With growth changes these fruit often appeared to be developing in the apical position. These patterns generally differed little from year to year. Variations may have been due to the differences in the number of flowers produced by the trees. Results were also similar between Hamlin and Valencia oranges. Patterns on Marsh grapefruit resembled those for the oranges but were frequently less consistent.     

Population ecology and phenology of Diaphorina citri (Hemiptera: Psyllidae) in two Florida citrus groves

Studies were conducted to assess population densities and phenology of the psyllid Diaphorina citri Kuwayama at two citrus groves in east-central Florida. One grove contained young, irrigated grapefruit trees and the other contained mature, nonirrigated orange trees. The two groves were sampled weekly for eggs, nymphs, and adults on flush shoots; for adults on mature leaves; and for adults captured on yellow sticky card traps. Because infestations of immature D. citri develop strictly on young flush, the abundance of flush was assessed weekly. Overall means of 26.5, 16.8, and 0.27 eggs, nymphs, and adults per flush shoot, respectively, were observed in the young grapefruit trees. In the grove of mature orange trees, overall means of 16.0, 12.7, and 0.31 eggs, nymphs, and adults per flush shoot were observed, respectively. Flush abundance was an inconsistent indicator of the mean density of D. citri per flush shoot. Mean density per shoot by itself was an inconsistent indicator of overall population levels of D. citri at each study site because few shoots were sometimes present when mean densities per shoot were high. May, June, and July were periods of time when immature D. citri were consistently present and most abundant at each study site, but the study indicated large infestations could occur at any time of the year depending on environmental factors and flush availability. Yellow sticky traps were effective for both male and female D. citri and useful for gauging adult population trends.     

Accumulation of Free Proline in Citrus Leaves during Cold Hardening of Young Trees in Controlled Temperature Regimes

Free proline increased in leaves of orange (Citrus sinensis [L.] Osb. cv. Valencia) and grapefruit (Citrus paradisi Macfad. cv. Star Ruby) trees on a wide range of citrus rootstocks during cold hardening. Increases in sugars accompanied proline accumulation. During cold hardening, the rate of proline accumulation was greater in old than in young leaves. In leaves of grapefruit trees kept in the dark during cold hardening, neither proline nor sugars increased and the degree of cold hardiness was less than in trees exposed to light. Like sugar accumulations, proline accumulation does not reflect specific degrees of cold hardiness in citrus cultivars.     

Developmental and physiological correlates of leaf size in Hyeronima alchorneoides (Euphorbiaceae)

The tropical emergent tree Hyeronima alchorneoides has large decreases in leaf size with tree age: 1200 cm(2) at 1 yr, 900 cm(2) at 3 yr, 200 cm(2) at 11 yr, and 80 cm(2) in old (>30 yr) individuals. We tracked leaf growth and physiological attributes on trees of three different ages (1, 3, and 11 yr) to determine the developmental basis and functional consequences of this variation. Leaves on young trees grew faster and sustained maximum rates of leaf expansion longer than leaves on older trees. Leaf mass per area (LMA) did not differ among age classes. Maximum photosynthetic rates reflected differences in leaf nitrogen concentration, in which leaves from the lower crown of younger trees outperformed those at a comparable crown position in older trees. One-year-old trees had the lowest stomatal conductance and the greatest instantaneous water use efficiency. Ontogenetic plasticity in mature leaf size, structure, and physiology may be a balance between the advantages conferred by rapid height growth when trees are young and the benefits derived from producing branches that increase light harvesting ability as trees reach the canopy.