Frost resistance and ice nucleation in leaves of five woody timberline species measured in situ during shoot expansion

Frost resistance and ice nucleation temperatures of leaves, from bud swelling until after full expansion, were measured in situ for five major woody timberline species with recently developed field freezing equipment. Frost resistance determined in situ on leaves of attached twigs was significantly higher than values determined on detached leaves in laboratory tests (e.g., the temperature at which incipient frost damage was observed (LTi) was 1.2 degrees C higher for detached leaves than for attached leaves of Picea abies (L.) Karst.). Frost resistance of leaves of all species changed significantly during shoot expansion (e.g., changes of 7.2 and 11 degrees C for Rhododendron ferrugineum L. and Larix decidua Mill., respectively). Expanding leaves (between 0 and 60% of full expansion) were the most sensitive to frost, with LTi values ranging from -3.4 degrees C in R. ferrugineum to -6.3 degrees C in L. decidua. Among the studied species, P. abies and R. ferrugineum were the most frost sensitive throughout the shoot elongation period. In situ freezing patterns of leaves of attached twigs also differed from those of leaves of excised twigs. During leaf expansion, two distinct freezing exotherms were always registered in situ. The first freezing event (E1, high-temperature exotherm) was recorded at -1.5 +/- 0.2 degrees C and reflected extracellular ice formation. Exposure of leaves to temperatures at which E1 occurred was, in all cases, noninjurious. The low-temperature exotherm (E2) mostly coincided with frost damage, except for some stages of leaf expansion in R. ferrugineum and P. abies, indicating that in situ freezing exotherms were not accurate estimators of frost damage in these species.     

Sources of N for leaf growth in a high-density apple (Malus domestica) orchard irrigated with ammonium nitrate solution

Elstar apple trees (Malus domestica Borkh.) on M.9 rootstock received either 5 or 35 g N tree(-1) year(-1) during the first two growing seasons after planting, applied as Ca(NO(3))(2) on a daily basis for nine weeks through a drip irrigation system. During the third growing season (1994), all trees were treated with 20 g N tree(-1) year(-1) as (15)NH(4) (15)NO(3) with applications starting on April 22 and continuing for 10 weeks. Soil solution nitrate-N and ammonium-N were monitored weekly with suction lysimeters located 30 cm beneath the drip emitters. Spur and shoot leaves were sampled intensively from full bloom to the end of rapid shoot growth. During the period of nitrogen application, soil solution nitrate-N and ammonium-N were relatively constant, at about 24 and 1.0 mg l(-1) respectively. Growth of the spur leaves was completed by one week after full bloom (May 12), whereas biomass of the shoot leaves increased until mid-June. Nitrogen for growth of the spur leaves was supplied mainly from remobilization, which was dependent on previous N supply. Accumulation of fertilizer N in spur leaves was independent of previous N treatments and continued until the end of the monitoring period (June 24), but contributed only 13% to total spur leaf N. Nitrogen for shoot leaf growth was independent of previous N treatments and was initially supplied primarily by remobilization, but by the end of extension growth, fertilizer N contributed 48% to total shoot leaf N. Linear increases in leaf N uptake throughout the period of rapid shoot growth and the large contribution of fertilizer N to total shoot leaf N were attributed to the constant supply of N available in the root zone through daily N fertilization.     

Translocation of nitrogen in the xylem of field-grown cherry and poplar trees during remobilization

Studies of small trees growing in pots have established that individual amino acids or amides are translocated in the xylem sap of a range of tree species following bud burst, as a consequence of nitrogen (N) remobilization from storage. This paper reports the first study of N translocation in the xylem of large, deciduous, field-grown trees during N remobilization in the spring. We applied 15N fertilizer to the soil around 10-year-old Prunus avium L. and Populus trichocharpa Torr. & Gray ex Hook var. Hastata (Dode) A. Henry x Populus balsamifera L. var. Michauxii (Dode) Farwell trees before bud burst to label N taken up by the roots. Recovery of unlabeled N in xylem sap and leaves was used to demonstrate that P. avium remobilizes N in both glutamine (Gln) and asparagine (Asn). Sap concentrations of both amides rose sharply after bud burst, peaking 14 days after bud burst for Gln, and remaining high some 45 days for Asn. There was no 15N enrichment of either amide until 21 days after bud burst. In the Populus trees, nearly all the N was translocated in the sap as Gln, the concentration of which peaked and then declined before the amide was enriched with 15N, 40 days after bud burst. Xylem sap of clonal P. avium trees was sampled at different positions in the crown to assess if the amino acid and amide composition of the sap varied within the crown. Sap was sampled during remobilization (when the concentration of Gln was maximal), at the end of remobilization and at the end of the experiment (68 days after bud burst). Although the date of sampling had a highly significant effect on sap composition, the effect of position of sampling was marginal. The results are discussed in relation to N translocation in adult trees and the possibility of measuring N remobilization by calculating the flux of N translocation in the xylem.     

Dry matter content during extension of twigs,buds and leaves reflects hydraulic status related to earlywood vessel development in <Emphasis Type="Italic">Quercus pyrenaica</Emphasis> Willd.

A quantitative method was tested to describe crown phenophases in relation to water content and to secondary growth in ring-porous species, based on the hypothesis that new shoots require hydrated tissues to maintain the necessary turgor for extension, leading to a reduction in dry matter content (DMC). We collected a three-year-old branch from 11 Quercus pyrenaica Willd. trees at 10-day intervals to estimate DMC of newly developing buds, leaves, and twigs, and processed two opposite stem microcores for xylogenesis. Branch phenophases and shoot length were recorded in the field. The DMC of all organs decreased during crown development, with a minimum in early June, followed by a gradual increase up to initial values in late September. The shoot extension period concurred with the lowest DMC, but also with the beginning of earlywood maturation in the main stem, suggesting a high tissue hydration only when earlywood vessels become functional to fulfill enough water requirements for shoot and leaf extension. These results confirm the usefulness of DMC to accurately quantify the phenology of primary growth from bud swelling up to full leaf extension, as a complement to qualitative methods. This variation in DMC appears to be linked to secondary growth as a result of earlywood vessel development.     

Stomatal conductance,growth and root signaling in Betula pendula seedlings subjected to partial soil drying

Seedlings of Betula pendula Roth were grown with their root systems separated between two soil compartments. Four treatments were imposed: (i) adequate irrigation in both compartments (WW, controls); (ii) adequate irrigation in one compartment and drought in the other compartment (WD); (iii) drought in both compartments (DD); and (iv) half of the root system severed and the remainder kept well-watered (root excision, RE). Predawn leaf water potential, stomatal conductance, soil-to-leaf specific hydraulic conductance, and root and leaf growth decreased in DD-treated seedlings, which also displayed severe leaf shedding (30% loss in leaf area). The DD treatment also resulted in increased concentrations of abscisic acid (ABA) and its glucose ester in the xylem sap of roots and shoots compared to concentrations in control seedlings (about 200 versus 20 nM). Despite the difference in xylem sap concentrations, total ABA flux to the shoots was similar in the two treatments (1-2 pmol ABA m(-2) leaf area s(-1)) as a result of reduced transpiration in the DD-treated seedlings. Compared with root growth in control plants, root growth increased in the RE-treated plants and decreased in the drying compartment of the WD treatment; however, the RE and WD treatments only slightly reduced leaf expansion, and had no detectable effects on shoot water relations or ABA concentrations of the root and shoot xylem sap. We conclude that short-term soil water depletion affecting only 50% of the root system does not cause a measurable stress response in birch shoots, despite root growth cessation in the fraction of drying soil.     

Effects of elevated CO(2) and temperature on cold hardiness and spring bud burst and growth in Douglas-fir (Pseudotsuga menziesii)

We examined effects of elevated CO(2) and temperature on cold hardiness and bud burst of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings. Two-year-old seedlings were grown for 2.5 years in semi-closed, sunlit chambers at either ambient or elevated (ambient + ~ 4 degrees C) air temperature in the presence of an ambient or elevated (ambient + ~ 200 ppm) CO(2) concentration. The elevated temperature treatment delayed needle cold hardening in the autumn and slowed dehardening in the spring. At maximum hardiness, trees in the elevated temperature treatment were less hardy by about 7 degrees C than trees in the ambient temperature treatment. In general, trees exposed to elevated CO(2) were slightly less hardy during hardening and dehardening than trees exposed to ambient CO(2). For trees in the elevated temperature treatments, date to 30% burst of branch terminal buds was advanced by about 6 and 15 days in the presence of elevated CO(2) and ambient CO(2), respectively. After bud burst started, however, the rate of increase in % bud burst was slower in the elevated temperature treatments than in the ambient temperature treatments. Time of bud burst was more synchronous and bud burst was completed within a shorter period in trees at ambient temperature (with and without elevated CO(2)) than in trees at elevated temperature. Exposure to elevated temperature reduced final % bud burst of both leader and branch terminal buds and reduced growth of the leader shoot. We conclude that climatic warming will influence the physiological processes of dormancy and cold hardiness development in Douglas-fir growing in the relatively mild temperate region of western Oregon, reducing bud burst and shoot growth.     

Terminal bud failure of black cottonwood (Populus trichocarpa) exposed to salt-laden winter storms

At coastal sites, trees are exposed to marine aerosols that may cause foliar necrosis and shoot dieback, which can result in deformed crowns and contorted stems. A six-year study of leaf primordia in terminal buds of black cottonwood trees (Populus trichocarpa Torr. & Gray) on Heimaey Island off the south coast of Iceland was undertaken to elucidate the physiological events associated with salt-deposition-related bud failure. Leaf and bud lengths, dry mass, water content and chloride concentrations were monitored and related to four phenological stages: (1) bud set; (2) dormancy induction; (3) dormancy release; and (4) bud break. The trees set buds in July and shed their leaves by late September. Leaf primordia generally stopped growing by September 10 +/- 22 days and attained midwinter water content in late September. Leaf growth commenced in the terminal buds by March 2 +/- 16 days, but mean dates of bud swelling and bud break were April 29 +/- 19 and May 10 +/- 12 days. In summer and until November, chloride concentrations in leaf primordia were low, but increasing. Chloride concentrations remained stable from December to February, even though the dormant trees were exposed to large amounts of marine aerosols. In February and March, three events occurred more or less simultaneously: (1) leaf extension growth commenced; (2) chloride concentration surged in the leaf primordia; and (3) the leaf primordia began to hydrate. Following dormancy release, growth and hydration of leaf primordia were negatively related to chloride concentration in the leaf primordia, with inhibition of leaf growth, tissue hydration and chloride acquisition occurring at a chloride concentration threshold estimated at 7.3 mg Cl- g(-1) tissue water. Necrosis of leaf primordia was observed above 14 mg Cl- g(-1) tissue water. Growth and hydration of leaves at bud break in mid-May was explained by a three-parameter logistic model of chloride concentration in leaf primordia at the end of March. By mid-May, 90% of all buds remained non-necrotic, but only 56% the terminal buds had broken. Salt alone explained the observed growth suppression of leaf primordia in the buds and the resultant failure of terminal buds to break by mid-May.     

Masting in Fagus crenata and its influence on the nitrogen content and dry mass of winter buds

In Fagus, full-mast seeding years are invariably followed by at least one non-mast year. Both flower and leaf primordia develop during the summer within the same winter buds. Flower bud initiation occurs when the N content of developing seeds is increasing rapidly. We hypothesized that competition for nitrogen (N) between developing seeds and buds limits flower primordium formation in mast years and, hence, limits seed production in years following mast years. We tested this hypothesis in three Fagus crenata Blume forests at elevations of 550, 900 and 1500 m. Bud N concentration (N con), amount of N per bud (N bud) and dry mass per bud (DM) were compared between a mast year (2005) and the following non-mast year (2006), and between winter buds containing both leaf and flower primoridia (BF), which were formed during the non-mast year, and winter buds containing leaf primordia only (BL), which were formed in both mast and non-mast years. In addition, leaf numbers per shoot corresponding to the analyzed buds were counted, and the effect of masting on litter production was analyzed by quantifying the amounts of litter that fell in the years 2004 to 2007. The dry mass and N content of BF formed in 2006 by trees at both 550 and 1500 m were 2.1-3.4-fold higher than the corresponding amounts in BL, although the numbers of leaves per current-year shoot in 2007 that developed from the two bud types in the same individuals did not differ significantly. These results indicate that more N and carbohydrate are expended in producing BF than in producing BL. The amount of litter from reproductive organs produced in the mast year was similar to the amount of leaf litter at 900 and 1500 m, but three times as much at 550 m. Leaf numbers per shoot were significantly lower at all elevations in the mast year than in the non-mast years (and the amount of leaf litter at 550 and 1500 m tended to be lower in the mast year than in the non-mast years. In conclusion, preferential allocation of resources to seeds in the mast year reduced the availability of resources for flower primordium formation, and this may have accounted for the poor seed production in the following non-mast year.     

Effects of winter buds on winter nitrogen uptake and allocation in Pinus densiflora saplings

Studies of nitrogen (N) use by plants have confirmed some winter N uptake; however, the mode of regulation of plant N use in winter is unknown. The regulation of N use by plants during winter may differ from that in the growing season, as plant growth strongly affects N use. We investigated the effects of winter buds on winter N use by Japanese red pine (Pinus densiflora), as a previous study demonstrated that N absorbed during winter contributes significantly to leaf growth in the following spring. We conducted a bud pruning experiment during winter to examine the effects of winter buds on winter N uptake and allocation among plant organs using ¹⁵N labeling. Over a three-week labeling period, the ¹⁵N content in roots increased to 0.20 ± 0.12 mg N g DW^?1, which is equivalent to 1.8 ± 1.1 % of the total N content in the roots. However, this absorbed ¹⁵N rarely appeared in needles and buds. Bud pruning did not affect ¹⁵N uptake and allocation. On the other hand, significant total N retranslocation was found within the crowns of saplings without bud pruning, but N was not retranslocated in bud-pruned plants. The bud pruning experiment indicated that N was retranslocated from needles into winter buds. Since soil N availability changes dramatically and is unstable in many forest ecosystems, N contained in needles would be a more stable source of N than newly absorbed N.     

Preformation in vegetative buds of Prunus persica: factors influencing number of leaf primordia in overwintering buds

We investigated the influence of bud position, cultivar, tree age, tree carbohydrate status, sampling date, drought and light exposure on the number of leaf primordia formed in dormant vegetative peach buds (Prunus persica (L.) Batsch) relative to the number of primordia formed after bud break (neoformed). During winter dormancy, vegetative peach buds from California and Italy were dissected and the number of leaf primordia recorded. Between leaf drop and bud break, the number of leaf primordia doubled from about five to about 10. Parent shoot length, number of nodes on the parent shoot, cross-sectional area of the parent shoot, bud position along the parent shoot and bud cross-sectional area were correlated with the number of leaf primordia. Previous season light exposure, drought and tree carbohydrate status did not affect the number of leaf primordia present. The number of leaf primordia differed significantly among peach varieties and tree ages at leaf drop, but not at bud break. Our results indicate that neoformation accounted for all shoot growth beyond about 10 nodes. The predominance of neoformed shoot growth in peach allows this species great plasticity in its response to current-season conditions.     

Seasonal changes in bud dormancy in relation to bud morphology, water and starch content, and abscisic acid concentration in adult trees of Betula pubescens

Annual cycles of change in bud morphology, bud burst ability, abscisic acid (ABA) concentration, and starch and water content were studied in mid-crown terminal buds of short shoots and underground basal buds of Betula pubescens Ehrh. In particular, we investigated the roles of ABA and bud water content in the regulation of bud growth. Basal buds differed morphologically from terminal buds of short shoots in that their leaf initials did not develop into embryonic foliage leaves and their total size did not increase significantly during summer. Bud burst ability, measured by forcing detached short shoots and stumps under controlled conditions, was maintained in the basal buds throughout the year, whereas the terminal buds of short shoots remained dormant until October, thereafter their bud burst ability increased gradually and reached a maximum in March-April. The ABA concentration of the basal buds was relatively constant throughout the sampling period (1-3 micro g g(DW) (-1)), whereas that of the terminal buds of short shoots, which was much higher (5-10 micro g g(DW) (-1)), showed a distinct seasonal cycle with a maximum from August to November. Bud ABA concentration decreased during the first 10 days of forcing, especially in basal buds. In both bud types, the amount of starch increased toward the autumn, declined in November, and was negligible in the terminal buds of short shoots between January and March, but in April, the amount was high again in both bud types. Water content varied characteristically in both bud types, although more distinctly in the terminal buds of short shoots, with an increase in spring before bud burst and a decrease during the summer until September. The significant morphological and physiological differences between the mid-crown terminal buds of short shoots and the underground basal buds may partly explain the characteristic growth habit of the basal buds and their development into coppice shoots after cutting the tree. The results also indicate a role for ABA in maintaining dormancy of the terminal buds of short shoots and emphasize the relationship between tissue water status and ABA concentration.     

库尔勒香梨树体生长与15N的吸收及分配动态

为给库尔勒香梨园合理施肥及氮肥利用率的提高提供参考,以6年生库尔勒香梨为研究对象,采用15N同位素示踪技术,研究萌芽前期至果实成熟期库尔勒香梨树体生长和氮素吸收、分配动态。结果表明:库尔勒香梨树体基径随着生育期的推移逐渐增大,于果实成熟期达到最大(8.71cm);库尔勒香梨叶片的叶面积指数、叶绿素SPAD值和叶片光合速率均随着香梨年生育期的推进呈现先增大后减小的趋势,均在第2个快速膨大期达到最大,分别为2.40、42.03和12.50μmol/(m^2·s);在年生育末期,库尔勒香梨单株树体的生物量为19958g,氮素积累量为199.44g,各器官中以当年新生器官果实的生物量和氮素积累量为最高,分别占整株树体生物量和氮素积累量的33.33%和25.08%。不同生育期15N在树体内的运转随生长中心的变化而变化。盛花期15N在1年生枝中的分配势最强,新梢旺长期和第2个快速膨大期15N在叶片中的分配势最强,果实成熟期15N在果实中的分配势最强。在果实成熟期库尔勒香梨树体当季15N肥料利用率为17.35%。     

Detection of the endophyte Discula umbrinella in buds and twigs of Fagus sylvatica

Endophytic fungi have been isolated from buds and twigs of beech trees collected at four different sites in Switzerland. Discula umbrinella. was recovered at high frequencies from the bud scales and the twig pieces contiguous to the buds, but was virtually absent from the rolled up leaves enclosed by the scales. In addition to infection by air-borne inoculum, thalli of D. umbrinella may grow from the twigs into the leaf tissues.     

Development and growth of primordial shoots in Norway spruce buds before visible bud burst in relation to time and temperature in the field

The timing of bud development in ecodormancy is critical for trees in boreal and temperate regions with seasonally alternating climates. The development of vegetative buds and the growth of primordial shoots (the primordial shoot ratio) in Norway spruce were followed by the naked eye and at stereo and light microscopic levels in fresh-cut and fixed buds obtained by regular field samplings during the spring of 2007, 2008 and 2009. Buds were collected from 15 randomly selected trees (all 16 years old in 2007) of one southern Finnish half-sib family. The air temperature was recorded hourly throughout the observation period. In 2008 and 2009, initial events in the buds, seen as accumulation of lipid droplets in the cortex area, started in mid-March and were depleted in late April, simultaneously with the early development of vascular tissue and primordial needles. In mid-April 2007, however, the development of the buds was at least 10 days ahead as a result of warm spells in March and early April. Variation in the timing of different developmental phases within and among the sample trees was negligible. There was no clear one-to-one correspondence between the externally visible and the internal development of the buds. The dependence of the primordial shoot ratio on different types of temperature sum was studied by means of regression analysis. High coefficients of determination (R(2)?≈?95%) were attained with several combinations of the starting time (beginning of the year/vernal equinox), the threshold value (from -3 to +5 °C), and the time step (hour/day) used in the temperature summation, i.e., the prediction power of the primordial shoot ratio models turned out to be high, but the parameter estimate values were not unambiguous. According to our results, temperature sums describe the growth of the primordial shoot inside the bud before bud burst. Thus, the results provide a realistic interpretation for the present phenological models of bud development that are based on temperature sums and external observations of bud burst only, and they also provide new tools for improving the models.     

Storage of foliar-absorbed nitrogen and remobilization for spring growth in young nectarine (Prunus persica var. nectarina) trees

The effectiveness of spraying foliage with urea to provide nitrogen (N) to augment the seasonal internal cycling of N in young nectarine trees (Prunus persica (L.) Batsch var. nectarina (Ait. f. Maxim.), cv. Stark Red Gold) was studied. One-year-old trees were grown with contrasting N supplies during the summer and foliage was sprayed with a 2% urea solution labeled with (15)N just before leaf senescence started. After leaf abscission had finished, the trees were repotted in sand and given no further N. Remobilization of both labeled and unlabeled N for leaf growth the following spring was quantified. Leaves absorbed between 58 and 69% of the (15)N intercepted by the canopy irrespective of tree N status. During leaf senescence, the majority of (15)N was withdrawn from the leaves into the shoot and roots. Remobilization of (15)N the following spring was also unaffected by tree N status. About 38-46% of (15)N in the trees was recovered in the new growth. More unlabeled N (derived from root uptake) was remobilized for leaf growth in the spring than was withdrawn from leaves during canopy senescence the previous autumn. Therefore, soil-applied N augmented N storage pools directly, and contributed more to N remobilization the following spring than did foliar-absorbed (15)N.     

Vigor-controlling rootstocks affect early shoot growth and leaf area development of kiwifruit

Patterns of shoot development and the production of different types of shoots were compared with scion leaf area index (LAI) to identify how eight clonal Actinidia rootstocks influence scion development. Rootstocks selected from seven Actinidia species (A. chrysantha Merri., A. deliciosa (A. Chev.) C. F. Liang et A.R. Ferguson, A. eriantha Benth., A. hemsleyana Dunn, A. kolomikta (Maxim. et Rupr.) Maxim., A. kolomikta C.F. Liang and A. polygama (Sieb. et Zucc.) Maxim.) were grafted with the scion Actinidia chinensis Planch. var. chinensis 'Hort16A' (yellow kiwifruit). Based on an earlier architectural analysis of A. chinensis, axillary shoot types produced by the scion were classified as short, medium or long. Short and medium shoots produced a restricted number of preformed leaves before the shoot apex ceased growth and aborted, resulting in a 'terminated' shoot. The apex of long shoots continued growth and produced more nodes throughout the growing seasons. Mid-season LAI of the scion was related to the proportion of shoots that ceased growth early in the season. Scions on low-vigor rootstocks had 50% or less leaf area than scions on the most vigorous rootstocks and had a higher proportion of short and medium shoots. On low-vigor rootstocks, a higher proportion of short shoots was retained during pruning to form the parent structure of the following year. Short parent shoots produced a higher proportion of short daughter shoots than long parent shoots, thus reinforcing the effect of the low-vigor rootstocks. However, overall effects of rootstock on shoot development were consistent regardless of parent shoot type and nodal position within the parent shoot. Slower-growing shoots were more likely to terminate and scions on low-vigor rootstocks produced a higher proportion of slow-growing shoots. Shoot termination also occurred earlier on low-vigor rootstocks. The slower growth of terminating shoots was detectable from about 20 days after bud burst. Removal of a proportion of shoots at the end of bud burst increased the growth rate and decreased the frequency of termination of the remaining shoots on all rootstocks, indicating that the fate of a shoot was linked to competitive interactions among shoots during initial growth immediately after bud burst. Rootstock influenced the process of shoot termination independently of its effect on final leaf size. Scions on low-vigor rootstocks had a higher proportion of short shoots and short shoots on all rootstocks had smaller final leaf sizes at equivalent nodes than medium or long shoots. Only later in the development of long shoots was final leaf size directly related to rootstock, with smaller leaves on low-vigor rootstocks. Thus, the most important effect of these Actinidia rootstocks on scion development occurred during the initial period of shoot growth immediately after bud burst.     

Nitrogen availability, local light regime and leaf rank effects on the amount and sources of N allocated within the foliage of young walnut (Juglans nigra x regia) trees

Early season leaf growth depends largely on nitrogen (N) provided by remobilization from storage, and many studies have tested the effect of N availability to roots on the amount of N provided for new leaf development by remobilization. Although it is well known that the light regime experienced by a leaf influences the amount of N per unit leaf area (LA), the effect of the local light regime on the amount of N derived either directly from root uptake or from remobilization for early season leaf growth has never been tested at an intra- canopy scale. The objective of this study was to quantify the relative importance of (1) N availability to roots, (2) local light regime experienced by the foliage (at the shoot scale) and (3) leaf rank along the shoot, on the total amount of N allocated to leaves and on the proportions of N provided by remobilization and root uptake. To quantify the importance of N uptake and remobilization as sources of leaf N, potted hybrid walnut trees (Juglans nigra L. x regia L.) were grown outdoors in sand and fed with a labeled ((15)N) nutrient solution. By removing the apical bud, the trees were manipulated to produce only two shoots. The experimental design had two factors: (1) high (HN; 8 mol N m(-3)) and low (LN; 2 mol N m(-3)) N availability; and (2) high (HL; 90% of incident photosynthetically active photon flux (PPF)) and low (LL; 10% of incident PPF) light. Total leaf N per tree was unaffected by either N availability or irradiance. The HN treatment increased the amount of leaf N derived from root uptake at the whole-tree scale (typically around 8 and 2% in the HN and LN treatments, respectively). Nitrogen allocation within foliage of individual trees was controlled by the local light regime, which strongly affected individual leaf characteristics as leaf mass per unit LA and area- based amount of leaf (N(a)). Decreasing the light availability to a branch decreased the amount of N allocated to it, benefiting the less shaded branches. In contrast, shading of the lower branch did not affect the fraction of total leaf N remobilized for either the lower, shaded branch or the upper, unshaded branch. The relevance of these findings for tree growth modeling is discussed.     

Shoot development,chlorophyll, gas exchange and carbohydrates in lychee seedlings (Litchi chinensis)

Shoot growth, chlorophyll concentrations, gas exchange and starch concentrations were studied in lychee (Litchi chinensis Sonn.) seedlings of cultivar "Wai Chee" grown in a heated greenhouse at Nambour in subtropical Australia (27 degrees S). We also examined the effects of shoot defoliation and root pruning on leaf expansion. Shoot growth showed a rhythmic cycle under constant greenhouse conditions, with a mean duration of flushing of 20 days and an interval of 10 days over three cycles. Shoots and leaves expanded in a sigmoidal pattern to about 80 mm and 500 cm(2), respectively, for each flush. Starch concentrations of the lower stem and roots decreased as the young red leaves expanded, and increased as the fully expanded leaves turned dark green. Chlorophyll concentrations and net CO(2) assimilation rate were highest in the fully expanded dark green leaves. Removing 50% of the area of each fully expanded leaf had little effect on the expansion of younger leaves, but total biomass of defoliated plants was only 60% of that of controls. In contrast, removing half the roots just before bud swelling reduced final leaf area by 80%. We conclude that the young shoot has relatively low rates of photoassimilation until the leaves are fully expanded and dark green, and depends on assimilates from elsewhere in the plant. During leaf expansion, translocation of assimilates to the shoot occurred at the expense of the roots.     

Environmental and physiological controls over oxygen and carbon isotope composition of Tasmanian blue gum, Eucalyptus globulus

We measured oxygen isotope ratios (delta18O) of xylem sap, phloem sap, leaves, wood and bark of Eucalyptus globulus Labill. growing in southwestern Australia. Carbon isotope ratios (delta13C) were measured in the dry matter of phloem sap, leaves and wood. Results were used to test several aspects of a mechanistic model of 18O enrichment and provided insights into post-photosynthetic variations in dry matter delta13C. Xylem water delta18O varied little within the tree crown, whereas variation at the landscape-level was more pronounced, with plantations near the coast being enriched by up to 3 per thousand compared with plantations less than 100 km inland. Phloem water was significantly enriched in 18O compared with xylem water in two of three sampling campaigns; mean enrichments were 0.5 and 0.8 per thousand. Phloem sap sugars exported from E. globulus leaves closely reflected observed leaf water enrichment when diurnal variation in photosynthesis was taken into account. Photosynthetic rates were higher in the morning than in the afternoon, whereas leaf water 18O enrichment increased to maximum values in the afternoon. A non-steady-state model of leaf water 18O enrichment accurately predicted observed values through a full diel cycle. Mean estimates of the proportion of organic oxygen effectively exchanging with xylem water during cellulose synthesis were close to 0.40 for both leaves and wood. Carbon isotope ratios of nascent xylem tissues did not differ from those of phloem sap sugars collected concurrently, whereas nascent leaf tissues were depleted in 13C by 2 per thousand compared with phloem sap sugars, suggesting that, in E. globulus, 13C enrichment of sink tissues compared with source leaves does not result from an enriching process within the sink tissue.