Mineral nutrition and leaf longevity in an evergreen shrub,Ledum palustre ssp. decumbens

Summary Effects of variable mineral nutrient status on evergreen leaf longevity were investigated in a field fertilization experiment, and by comparison of plants from several sites. The species studied was Ledum palustre spp. decumbens, with a normal leaf life expectancy of 2.06 years and a maximum leaf longevity of 4.5–5.0 years. Most leaf losses took place during the growing season, not during the winter. Fertilization increased leaf production but decreased leaf survivorship. Total number of leaves per stem was unchanged with fertilization. In a comparison among sites, there was a moderate negative correlation between plant N and P concentrations and leaf longevity. These intraspecific responses are similar to known interspecific changes along nutrient gradients, i.e. with high nutrient availability a vegetation should become more deciduous and less evergreen.     

Plant growth strategies in evergreen-sclerophyllous shrublands (Maquis) in central Greece

A synchronic method was applied in order to investigate the growth process of the dominant woody plants in an undisturbed Mediterranean evergreen-sclerophyllous ecosystem in central Greece. Cover (%), biomass and leaf area data of Arbutus unedo, Erica arborea, Quercus coccifera and Phyllyrea latifolia over a period of 30 years are provided. The investigation of growth by means of the Richards function revealed two plant groups, following two different growth strategies: A fast growth strategy, followed by A. unedo and E. arborea and a slow growth strategy followed by Q. coccifera and P. latifolia. Furthermore, a succession pattern was described in which the fast growing initial dominants are being replaced by the slow growing plants. Two succession scenaria are proposed as alternative plant responses to grazing and fires, which are the major anthropogenic impacts in the mediterranean environment.     

Assessment of within-population genetic structure in <Emphasis Type="Italic">Quercus crispula</Emphasis> and <Emphasis Type="Italic">Q. dentata</Emphasis> by amplified fragment length polymorphism analysis

The spatial genetic structure was assessed by amplified fragment length polymorphism (AFLP) technique for Quercus crispula Blume (37 individuals) and Q.dentata Thunberg (54 individuals) growing along a 550-m transect in Ishikari, Hokkaido, northern Japan. The simple Mantel test revealed significant negative correlation between genetic similarity and geographic distance in Q.dentata and negative but insignificant correlation in Q.crispula. Spatial autocorrelation analysis revealed that Mantels r generally decreased from positive to negative values with the increase of spatial distance in both oak species with significant deviation from zero for the 50–100-m (positive) and 250–300-m classes (negative) in Q.dentata. Thus, significant spatial genetic structure was revealed at least in Q.dentata     

Seasonal variation in leaf chemistry of the coast live oak Quercus agrifolia and implications for the California oak moth Phryganidia californica

Summary The perennial foliage of the California coast live oak (Quercus agrifolia Nee) permits herbivores to feed on this oak species throughout the year. Patterns of herbivory for a two-year period on Q. agrifolia were observed in relation to seasonal and age-related changes in the nutritional and defensive characteristics of leaves. Nitrogen and phosphorus contents were higher in new leaves compared to mature foliage. Structural compounds (e.g., cellulose) in leaves rapidly increased with age. Concentrations of tatal phenolics (Folin-Denis) and astringency were higher in new foliage, and concentrations of condensed tannins gradually increased as the leaves matured. Peaks of herbivore damage were observed in June and in September–October, and were caused by outbreaks of the California oak moth (Phryganidia californica). P. californica, a bivoltine oak specialist, exhibited feeding preferences in June for old leaves over emerging leaves, and showed no preferences for leaf classes in September. These results suggest that P. californica is adapted to survive on nutritionally poor foliage and to circumvent quantitative defenses such as condensed tannins.     

Limitations due to water stress on leaf net photosynthesis of Quercus coccifera in the Portuguese evergreen scrub

Summary Gas exchange characteristics in leaves of the sclerophyll shrub Quercus coccifera were studied in the natural habitat in Portugal during spring and during the summer dry period. Compared to other sclerophyll species growing at the same site, photosynthesis in leaves of Quercus coccifera was less affected by water stress. Moderate water stress after six weeks of drought led to large decreases in stomatal conductance but no change in mesophyll photosynthetic capacity as compared to late spring. Leaf internal CO₂ pressure remained near 220 bar during diurnal courses in the spring. On midsummer days, leaf internal CO₂ decreased from a late morning value of 200 bar to a late afternoon value of approximately 150 bar. In contrast to Quercus suber (Tenhunen et al. 1984), restriction of CO₂ supply due to stomatal closure reduced net CO₂ uptake at midday and in the afternoon during midsummer. A decrease in leaf carboxylation efficiency and an increase in CO₂ compensation point at midday also played an important role in determining the diurnal course of net photosynthesis. During the late stages of drought in September, severe water stress led to reduction in mesophyll photosynthetic capacity and further reduction in leaf conductance. The observed decrease in mesophyll photosynthetic capacity was correlated with decrease in the daily minimum leaf water potential to greater negative values than-30 bar. At this time, CO₂ saturated photosynthetic rates decreased as much as 50% over the course of a day when measured at constant saturating light, 32° C leaf temperature, and a water vapor mole fraction difference between leaf and air of 30 mbar bar^-1.     

Feeny revisited: condensed tannins as anti-herbivore defences in leaf-chewing herbivore communities of Quercus

Abstract. 1. Community level oak–tannin–insect patterns have been largely unexplored since Paul Feeny's ground‐breaking research. Two hypotheses were tested for Quercus velutina and Q. alba in the Missouri Ozarks: abundance and richness of leaf‐chewing herbivores are negatively correlated with foliar condensed tannin concentrations and variation in condensed tannin concentrations explains variation in herbivore community structure. 2. In 2001, foliar condensed tannins in the understorey and canopy of these two oak species were quantified simultaneously with censuses of herbivores in May, during leaf expansion, and in June and August, when leaves were fully expanded. Thirty‐eight of the 134 species encountered had densities sufficient to be analysed individually (n = 10). Of those, Acronicta increta (Noctuidae) and Attelabus sp. (Curculionidae), both oak specialists, were negatively correlated with condensed tannins in the canopy of Q. alba. One additional specialist, Chionodes pereyra (Gelechiidae), was marginally negatively correlated with condensed tannins in the understorey of Q. velutina. Understorey species richness of May Q. velutina herbivores was negatively correlated with condensed tannins, as were total canopy insect density and species richness of August herbivores on Q. alba. 3. Principal component analysis (PCA) of insect abundances indicated that understorey and canopy Q. velutina and Q. alba had different communities of leaf‐chewing insects. Furthermore, condensed tannin levels contributed significantly to variation in PCA scores for Q. velutina, explaining 25% of the total variation. 4. Overall, these results indicate that specialists were more likely than generalists both to correlate negatively with condensed tannins and to occur in lower tannin habitats; abundance and richness of both early and late season fauna correlated negatively with tannins; and species were more likely to correlate negatively with condensed tannins when feeding on Q. alba than on Q. velutina and when feeding in the canopy than in the understorey. Future studies of tannin–insect interactions should manipulate leaf quality in combination with manipulations of other factors that likely influence community structure.     

Carbon isotope composition in relation to leaf gas exchange and environmental conditions in Hawaiian Metrosideros polymorpha populations

Summary Carbon isotope composition, photosynthetic gas exchange, and nitrogen content were measured in leaves of three varieties of Metrosideros polymorpha growing in sites presenting a variety of precipitation, temperature and edaphic regimes. The eight populations studied could be divided into two groups on the basis of their mean foliar ¹³C values, one group consisting of three populations with mean ¹³C values ca.-26 and another group with ¹³C values ca.-28. Less negative ¹³C values appeared to be associated with reduced physiological availability of soil moisture resulting from hypoxic conditions at a poorly drained high elevation bog site and from low precipitation at a welldrained, low elevation leeward site. Gas exchange measurements indicated that foliar ¹³C and intrinsic wateruse efficiency were positively correlated. Maximum photosynthetic rates were nearly constant while maximum stomatal conductance varied substantially in individuals with foliar ¹³C ranging from-29 to-24. In contrast with the patterns of ¹³C observed, leaf nitrogen content appeared to be genetically determined and independent of site characteristics. Photosynthetic nitrogenuse efficiency was nearly constant over the range of ¹³C observed, suggesting that a compromise between intrinsic water- and N-use efficiency did not occur. In one population variations in foliar ¹³C and gas exchange with leaf cohort age, caused the ratio of intercellular to atmospheric partial pressure of CO₂ predicted from gas exchange and that calculated from ¹³C to be in close agreement only in the two youngest cohorts of fully expanded leaves. The results indicated that with suitable precautions concerning measurement protocol, foliar ¹³C and gas exchange measurements were reliable indicators of potential resource use efficiency by M. polymorpha along environmental gradients.     

The effect of long term CO2 enrichment on the growth,biomass partitioning and mineral nutrition of Sitka spruce (Picea sitchensis (Bong.) Carr.)

Sitka spruce [Picea sitchensis (Bong.) Carr.] seedlings were grown for 3 years in an outside control plot or in ambient (355 mol mol^-1) or elevated (ambient + 350 mol mol^-1) atmospheric CO₂ environments, within open top chambers (OTCs) at the Institute of Terrestrial Ecology, Edinburgh. Sequential harvests were carried out at the end of each growing season and throughout the 1991 growing season, five in all. Plants grown in elevated CO₂ had, (i) 35 and 10% larger root/shoot ratios at the end of the first and third season, respectively, (ii) significantly higher summer leader extension relative growth rates, which declined more rapidly in early autumn than ambient grown plants, (iii) after three growing seasons a significantly increased mean annual relative growth rate, (iv) consistently lower foliar nutrient concentrations, and (v) after two growing seasons smaller total projected needle areas. Plants grown inside OTCs were taller, heavier and had a smaller root/shoot ratio than those grown outside the chambers. There was no effect of CO₂ concentration on Sitka spruce leaf characteristics, although leaf area ratio, specific leaf area and leaf weight ratio all fell throughout the course of the 3 year experiment.     

Comparative water relations of four Mediterranean oak species

The water relations and responses of two evergreen (Quercus ilex L. and Q. suber L.) and two deciduous (S. afares Pomel. and Q. faginea Will.) Quercus species were studied under experimental conditions. Two-year old seedlings grown in 30 l. pots were subjected to a drying period during which stomatal conductance, pre-dawn potential and minimum foliar potential were measured.The results shows that, for all species, the daily course of stomatal conductance agrees with the patterns proposed by Hinckley et al. (1978 & 1983). Concurrent with the species responses to short-term variation in water availability, it was found that pre-dawn leaf water potential controlled the maximum daily leaf conductance. There was a strong correlation between pre-dawn leaf potential and maximum daily conductance as described by the reciprocal function g_{srmax for}=(-0.47+2.61._p)^-1 the evergreen oaks and g_{srmax for}=(-1.94+7.39._p)^-1 for the deciduous species. These differences between the two groups may partialy explain their geograhic distributions, and suggest general questions concerning the mechanisms which optimize water-use efficiency in Mediterranean oak species.     

Polyphenolics in <Emphasis Type="Italic">Rhizophora mangle</Emphasis> L. leaves and their changes during leaf development and senescence

The chemical defenses in Rhizophora mangle L. are largely carbon based. The family has long been exploited for the high proanthocyanidin (condensed tannin) content of its wood, bark and leaves. In this paper, we quantify the overall pools of plant phenolics in R. mangle leaves, identify the major constituents of these pools and document their changes during leaf maturation and senescence. Overall, polyphenolics account for approximately 23% of the total leaf dry weight. The leaves contain at least seven flavonoid glycosides, five of them based on quercetin. Additional minor constituents are myricetin and kaempferol diglucosides. The aglycone, quercetin, was found only in senescing leaves. Also during senescence, a new compound, 5,4-dimethoxy-7,3,5-trihydroxyflavone, appeared. The flavonoids were accompanied by a complex mixture of condensed tannins based mainly on (+)-catechin and (–)-epicatechin with A-type and B-type linkages; this pool is also distinguished by having previously unreported, high contributions of (+)-catechin and (–)-epicatechin glycosides. During senescence, but prior to leaf abscission, the polyphenolic pools become simplified: flavonol glycosides and low oligomeric tannins largely disappear, leaving only the largest tannin polymers. The ecological and physiological significance of these compounds as they appear in R. mangle is discussed.     

Effects of short-term storage of gametes on fertilization of Pacific herring eggs

A method is described whereby arrays of samples ofClupea pallasi eggs may be stored during their preparation. The high fertilization potential retained by the eggs during short-term storage allows them to be fertilized synchronously when sample preparation is complete. A variation of the dry method of storage retained maximum fertilization potential (80–85%) of the eggs for about 1 hr, and of milt dilution (18 with 17 S sea water), about 7 hr. Following dry storage, eggs fertilized in salinities of 0–45 showed maximum rates of fertilization in salinities of 10–20, and fertilization rates 50% in salinities of 4.5–42. Salinities of fertilization influenced egg diameter, median hatching time, and larval length at hatching in egg samples transferred 21/2 hr after fertilization to an incubation salinity of 17 at 7°C. Fertilization rates were higher (90–95%) for eggs stored in 17 S at 7°C prior to fertilization. Under such wet storage conditions, maximum fertilization pontential was retained for about 2 hr. Highest fertilization rates (95–96%) were obtained for eggs stored and fertilized in salinities of 12–15. For the species and the area of origin considered (British Columbia), wet storage of eggs should result in maximum fertilization when the eggs are stored at 4°C for a period not greater than 2 hr prior to fertilization in the 12–15 S storage medium.Prepared under the auspices of the Canadian-German Scientific and Technical Cooperation Agreement.     

Genetic modification of condensed tannin biosynthesis in Lotus corniculatus. 1. Heterologous antisense dihydroflavonol reductase down-regulates tannin accumulation in “hairy root” cultures

An antisense dihydroflavonol reductase (DFR) gene-construct made using the cDNA for DFR from Antirrhinum majus was introduced into the genome of a series of clonal genotypes of Lotus corniculatus via Agrobacterium rhizogenes. After initial screening, 17 antisense and 11 control transformation events were analysed and tannin levels found to be reduced in antisense root cultures. The effect of this antisense construct, (pMAJ2), which consisted of the 5 half of the DFR cDNA sequence, was compared in three different recipient Lotus genotypes. This construct effectively down-regulated tannin biosynthesis in two of the recepient genotypes (s33 and s50); however, this construct was relatively ineffective in a third genotype (s41) which accumulated high levels of condensed tannins in derived transgenic root cultures. Four pMAJ2 antisense and three control lines derived from clonal genotypes s33 and s50 were selected and studied in greater detail. The antisense DFR construct was found to be integrated into the genome of the antisense hairy root cultures, and the antisense RNA was shown to be expressed. Tannin levels were much lower in antisense roots compared to the controls and this reduction in tannin levels was accompanied by a change in condensed tannin subunit composition.     

Water relations and growth of shrubs before and after fire in a semi-arid woodland

Summary Plant water relations and shoot growth rate of shrubs resprouting after fire or unburnt were measured in a semi-arid poplar box (Eucalyptus populnea) shrub woodland of eastern Australia. In vegetation unburnt for about 60 years, the dawn xylem water potential (_x) of the dominant shrub species was about-1.0 MPa when the soil was wet and-8.0 MPa when the soil was very dry. At any one time, the dominant shrub species,Eremophila mitchellii, E. sturtii, Geijera parviflora andCassia nemophila, were similar in _x butAcacia aneura andDodonaea viscosa were consistently higher in _x than this group when the soil was moist and lower when the soil was dry. The dominant tree species,Eucalyptus populnea andE. intertexta, appeared to have access to additional water beneath the hardpan which is located 60–80 cm below the surface. When shrubs were under extreme water stress (_x of-8 MPa), the trees had a _x of-3 to-3.6 MPa. Following a fire, both _x and leaf stomatal conductance (g _s) of resprouting shrubs were higher for about 5 years than comparable-aged unburnt vegetation, with relative differences in _x increasing with drought stress. Elongation rate of resprouts was positively linked to prefire shrub height in 3 of 4 species. However, shrubs resprouting after high intensity fires had substantially higher rates of shoot elongation than after low intensity fires which were in turn higher than for foliar expansion of unburnt shrubs. It is concluded that the growth rate of resprouting shrubs is primarily determined by physiological/ morphological factors associated with plant size but is also assisted by greater availability of water and possibly nutrients for a period after fire.     

Fine root biomass,production and turnover in a fertilized <Emphasis Type="Italic">Larix leptolepis</Emphasis> plantation in central Korea

The effects of fertilization [control (C), 200kgNha^–1+25kgP ha^–1 (LNP) and 400kgNha^–1+ 50kgP ha^–1 (HNP)] on fine root dynamics were examined in a 40-year-old Larix leptolepis plantation in central Korea. The average fine root biomass during the growing season for C, LNP and HNP was 957, 934 and 814kgha^–1, respectively, whereas the fine root production for C, LNP and HNP was 2103, 2131 and 2066kgha^–1, respectively. Nitrogen and P inputs into the soil via fine root turnover for C, LNP and HNP were 23.0 and 1.2, 23.3 and 1.2 and 22.6 and 1.2kgha^–1, respectively. There were no significant differences in fine root biomass, production and N and P inputs through fine root turnover between the fertilization treatments during the first growing season after fertilization.     

Ecophysiological analysis of woody species in contrasting temperate communities during wet and dry years

This study employed an intensive sampling regime in which leaf gas exchange and tissue-water relations were measured simultaneously on the same leaf at midday on 19 tree species from three distinct forest communities during wet (1990) and dry (1991) growing seasons. The study sites were located on a xeric barrens, a misic valley floor, and a wet-mesic floodplain in central Pennsylvania, United States. The xeric, mesic, and wetmesic sties had drought-related decreases in gravimetric soil moisture of 53, 34 and 27%, respectively. During the wet year, xeric and mesic communities had high seasonal mean photosynthetic rates (A) and stomatal conductance of water vapor (g _wv) and low midday leaf water potential (), whereas the wet-mesic community had low A and g _wv and high midday . The mesic and wet-mesic communities had dry year decreases in predawn , g _wv and A with the greatest drought effect occurring in the mesic community. Regression analysis indicated that species from each site that exhibited high wet-year A and g _wv tended to have low midday . This trend was reversed only in the mesic community in the drought year. Despite differences in midday , all three communities had similar midday leaf turgor pressure (_p) in the wet year attributable to lower osmotic potential at zero turgor ( ⁰ ) with increasing site droughtiness. Lower wet year ⁰ in the xeric community was due to low symplast volume rather than high solute content. Species with the lowest ⁰ in the wet year often did not have the lowest ¹⁰⁰ possibly related to differences in tissue elasticity. Moreover, increased elasticity during drought may have masked osmotic adjustment in ¹⁰⁰ but not in ⁰ , via dilution of solutes at full hydration in some species. Despite the sampling regime used, there were no relationships between gas exchange and osmotic and elastic parameters that were consistently significant among communities or years. This result questions the universal, direct effect of osmotic and elastic adjustments in the maintenance of photosynthesis during drought. By including a large number of species, this study provided new insight to the ecophysiology of contrasting forest communities, and the community-wide impact of drought on contrasting sites.     

Competition between <Emphasis Type="Italic">Eucalyptus victrix</Emphasis> seedlings and grass species

Competition in a natural system may be interspecific or intraspecific. In semiarid ecosystems, competition for resources between established neighboring grass species and newly recruited seedlings is very high. To examine the effects of grass species density, growing space and time of establishment on Eucalyptus victrix seedlings (interspecific competition), and the effect of density and growing space within E.victrix (intraspecific competition) we conducted an experiment under controlled conditions. We tested four hypotheses (i) E.victrix seedling growth is not affected by grass density; (ii) there is no difference in E.victrix survival and growth between early and later grass establishment; (iii) interspecific competition is not more intense than intraspecific competition in E.victrix; and (iv) growth of E.victrix seedlings is not dependent on available growing space. In a monoculture of E.victrix, seedling mortality was higher (10%) in large pots. In mixed culture pots, where E.victrix seedlings and grass seedlings were planted on the same day, E.victrix seedlings survived for up to 4weeks, but started to die after week five in the smallest pots. However, mortalities occurred in pots of all sizes when grass was established before E.victrix seedlings. Results also indicated that the resources necessary for the growth of individual E.victrix seedlings were more limiting under conditions of increased density of neighboring grass species rather than intraspecific competition. In particular, photosynthetic area of E.victrix seedlings was drastically reduced in mixed cultures. Although density, pot size and time of planting had impacts on E.victrix seedlings, the patterns of these impacts were variable.     

Stipularbildungen beiDentaria und ihr Wert für die morphologische Deutung der Speicherschuppen als Phyllodien

Zusammenfassung An den schuppenförmigen, speichernden Niederblättern vonDentaria enneaphyllos L.,D. bipinnata C. A.Mey.,D. pentaphyllos L. undDentaria bulbifera L. sowie an den Schuppenblättern der Bulbillen der letztgenannten Art wurden Bildungen aufgefunden, die in Form, Stellung und Entstehung den Nebenblättern an den Laubblättern gleichen und auch als solche aufgefaßt werden müssen. Aus der basalen Insertion dieser Stipeln und aus entwicklungsgeschichtlichen Befunden ergibt sich, daß das Unterblatt am Aufbau des Niederblattes nur äußerst geringen Anteil hat. Die Schuppenblätter müssen daher nicht wie bisher mit dem Blattgrund, sondern mit einem Teil des Oberblattes — nämlich dem Blattstiel (die Spreite ist nur als Rudiment vertreten) homologisiert werden und können somit als Phyllodien gelten. Um den Niederblattcharakter zu betonen, wird man zweckmäßigerweise den Ausdruck Blattstiel- oder Phyllodialschuppen verwenden. Diese Blattstielschuppen stellen eine besondere und bisher unbekannte Gestaltungsmöglichkeit speichernder Niederblätter dar.

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Stipules inDentaria and their importance for the morphological interpretation of the storage scales as phyllodes

Summary Cruciferae are commonly said to have no stipules, but some authors have correctly pointed out that tiny stipules are nearly always present and form a regular morphological component of the leaf structure in this family (cf.Weberling, 1956).The present investigations refer to the stipules and the morphology of the storage scales on the rhizomes inDentaria enneaphyllos L.,D. bipinnata C. A.Mey.,D. pentaphyllos L.,D. bulbifera L., and on the bulbils of the last mentioned species. These cataphylls have usually and up to now been interpretated as corresponding to the leaf base (in the sense of the German Unterblatt), the leaf stalk and lamina being rudimentary. The present results correct this conception: Ontogeny shows that an early articulation into leaf base (carrying the stipules) and upper leaf (Oberblatt) does take place, but the leaf base completes its development very early, being restricted therefore in the adult cataphyll to a narrow basal zone, the tiny stipules being placed quite near the attachment of the scale. Hence, the cataphyll is not to be identified with the leaf base, but is homologous to the leaf stalk with a rudimentary lamina. Therefore it can be called a phyllode. For a more precise morphological terminology the terms leaf stalk scale or phyllodial scale might be applicable. These phyllodial scales represent a hitherto unknown type of morphological organization among storage cataphylls.

     

Effect of nitrogen fertilization upon the secondary chemistry and nutritional value of quaking aspen (Populus tremuloides Michx.) leaves for the large aspen tortrix (Choristoneura conflictana (Walker))

Summary We investigated the effects of nitrogen fertilization upon the concentrations of nitrogen, condensed tannin and phenolic glycosides of young quaking aspen (Populus tremuloides) leaves and the quality of these leaves as food for larvae of the large aspen tortrix (Choristoneura conflictana), a Lepidopteran that periodically defoliates quaking aspen growing in North America. Nitrogen fertilization resulted in decreased concentrations of condensed tannin and phenolic glycosides in aspen leaves and an increase in their nitrogen concentration and value as food for the large aspen tortrix. These results indicate that plant carbon/nutrient balance influences the quality of aspen leaves as food for the large aspen tortrix in two ways, by increasing the concentrations of positive factors (e.g. nitrogen) and decreasing the concentrations of negative factors (eg. carbon-based secondary metabolites) in leaves. Addition of purified aspen leaf condensed tannin and a methanol extract of young aspen leaves that contained condensed tannin and phenolic glycosides to artificial diets at high and low levels of dietary nitrogen supported this hypothesis. Increasing dietary nitrogen increased larval growth whereas increasing the concentrations of condensed tannin and phenolic glycosides decreased growth. Additionally, the methanol extract prevented pupation. These results indicate that future studies of woody plant/insect defoliator interactions must consider plant carbon/nutrient balance as a potentially important control over the nutritional value of foliage for insect herbivores.     

Pigment dynamics and autumn leaf senescence in a New England deciduous forest,eastern USA

The leaves of woody plants at Harvard Forest in Central Massachusetts, USA, changed color during senescence; 70% (62/89) of the woody species examined anatomically contained anthocyanins during senescence. Anthocyanins were not present in summer green leaves, and appeared primarily in the vacuoles of palisade parenchyma cells. Yellow coloration was a result of the unmasking of xanthophyll pigments in senescing chloroplasts. In nine red-senescing species, anthocyanins were not detectable in mature leaves, and were synthesized de novo in senescence, with less than 20µg cm^–2 of chlorophyll remaining. Xanthophyll concentrations declined in relation to chlorophyll to the same extent in both yellow- and red-leaved taxa. Declines in the maximum photosystemII quantum yield of leaves collected prior to dawn were only slightly less in the red-senescing species, indicating no long-term protective activity. Red-leaved species had significantly greater mass/area and lower chlorophylla/b ratios during senescence. Nitrogen tissue concentrations in mature and senescent leaves negatively correlated to anthocyanin concentrations in senescent leaves, weak evidence for more efficient nitrogen resorption in anthocyanic species. Shading retarded both chlorophyll loss and anthocyanin production in Cornus alternifolia, Acer rubrum, Acer saccharum, Quercus rubra and Viburnum alnifolium. It promoted chlorophyll loss in yellow-senescing Fagus grandifolia. A reduced red:far-red ratio did not affect this process. Anthocyanins did not increase leaf temperatures in Q.rubra and Vaccinium corymbosum on cold and sunny days. The timing of leaf-fall was remarkably constant from year to year, and the order of senescence of individual species was consistent.     

Pioneer and late stage tropical rainforest tree species (French Guiana) growing under common conditions differ in leaf gas exchange regulation,carbon isotope discrimination and leaf water potential

Leaf gas exchange rates, predawn _wp and daily minimum _wm leaf water potentials were measured during a wet-to-dry season transition in pioneer (Jacaranda copaia, Goupia glabra andCarapa guianensis) and late stage rainforest tree species (Dicorynia guianensis andEperua falcata) growing in common conditions in artificial stands in French Guiana. Carbon isotope discrimination () was assessed by measuring the stable carbon isotope composition of the cellulose fraction of wood cores. The values were 2.7 higher in the pioneer species than in the late stage species. The calculated time integratedC _i values derived from the values averaged 281 mol mol^–1 in the pioneers and 240 mol mol^–1 in the late stage species. The corresponding time-integrated values of intrinsinc water-use efficiency [ratio CO₂ assimilation rate (A)/leaf conductance (g)] ranged from 37 to 47 mmol mol^–1 in the pioneers and the values were 64 and 74 mmol mol^–1 for the two late stage species. The high values were associated—at least inJ. copaia—with high maximumg values and with high plant intrinsinc specific hydraulic conductance [Cg/(_wm–_wp], which could reflect a high competitive ability for water and nutrient uptake in the absence of soil drought in the pioneers. A further clear discriminating trait of the pioneer species was the very sensitive stomatal response to drought in the soil, which might be associated with a high vulnerability to cavitation in these species. From a methodological point of view, the results show the relevance of for distinguishing ecophysiological functional types among rainforest trees.