SHOOT GROWTH AND HISTOGENESIS OF TREES POSSESSING DIVERSE PATTERNS OF SHOOT DEVELOPMENT

Shoot growth and histogenesis were followed in five unrelated tree taxa possessing inherently diverse patterns of shoot development. Following the resumption of growth in spring, each species differs quantitatively in the number of internodes elongating contemporaneously, in rates and duration of internodal elongation and seasonal periodicity of shoot growth. The basic pattern of internode elongation and histogenesis is qualitatively similar in each of the dicotyledonous species observed irrespective of growth habit or final form of the shoot produced. During the intial phase of internode development, growth is essentially uniform throughout young internodes, corresponding to an active period of cell division during which time pith cells increase in size to about one-third their final length. Subsequently, the pattern of cell division shifts progressively upward concomitant with increased elongation and maturation of pith cells in the basal portion of developing internodes. Thereafter, a wave of cell division accompanied by cell elongation continues to proceed acropetally until growth finally ceases in the distal portion of each internode. As long as internode elongation continues, frequently at distances 15–20 cm below the shoot apex, cell divisions still occur in the distal growing portion. As successive portions of each internode mature acropetally, final length of pith cells becomes relatively uniform throughout the internode. During the process of internode growth and development, cell lengths increase only two- to threefold, whereas cell numbers increase ten- to 30-fold, indicating the dominant role of cell division and increases in cell number to final internode length. Morphological patterns of shoot expression associated with differences in internode lengths along the axis of either preformed or neoformed shoots, as well as sylleptic branches, are due to differences in cell number rather than final cell length. Significant variations in final internode lengths along the axis of episodic shoots, caused by either endogenous or exogenous factors, are also attributed to differences in cell number.     

Clarifying the effects of dwarfing rootstock on vegetative and reproductive growth during tree development: a study on apple trees

BACKGROUND AND AIMS: Despite the widespread use of dwarfing rootstocks in the fruit-tree industry, their impact on tree architectural development and possible role in the within-tree balance between growth and flowering are still poorly understood, in particular during the early years of growth. The present study addressed this question in apple trees, through a detailed analysis of shoot populations, i.e. both vegetative and flowering shoots, during tree development. METHODS: Architectural databases were constructed for trees of two cultivars that were either own-rooted or grafted on dwarfing rootstock. Within-tree shoot demographics and annual shoot characteristics, i.e. their dimensions, number of laterals and flowering, were observed from the first to the fifth year of growth and compared among scion/root system combinations. KEY RESULTS: Differences in axis demographics appeared among scion/root system combinations after the second year of growth. Differences were found (a) in the number of long axes and (b) the number of medium axes. Dwarfing rootstock reduced the total number of axes developed in a tree, and this reduction resulted from proportionally more medium axes and spurs than long axes. The life span of spurs was also shortened. These phenomena appeared after an increase in flowering that started in the second year of growth and involved both axillary and terminal positions. Flowering regularity was also increased in grafted trees. CONCLUSIONS: These results confirm that the number of long shoots and flowering potential depend on the cultivar. They indicate that tree architectural plasticity in response to its root system mainly derives from the number of medium shoots developed and follows priorities within the whole tree axis population. There was also evidence for dwarfing rootstock involvement in adjusting the flowering abundance and that differences in flowering occurrence take precedence over those regarding vegetative growth during tree development.     

Growth Dynamics and Morphology of Annual Shoots, According to their Architectural Position, in Young Cedrus atlantica (Endl.) Manetti ex Carriere (Pinaceae)

The influence of shoot architectural position on growth andbranching pattern of young Cedrus atlantica (Endl.) Manettiex Carrière trees were studied. Extension growth andtype of axillary products (lateral bud, sylleptic short or longshoots) of annual shoots of increasing branching order (mainstem, branches and branchlets) were recorded weekly during the1993 growing season. Annual final shoot length, duration ofextension, and maximum extension rate decreased with increasingbranching order. Sylleptic axillary shoots occurred only onannual shoots of the main stem and branches and were producedwhen extension rate was at its highest. Differences in growthrate and final length of annual shoots, according to their architecturalposition, were related to differences in the total number anddiversity of types of sylleptic axillary shoots produced. Itis suggested that types and numbers of sylleptic axillary shootsproduced are linked with threshold values for both final lengthand extension rate of the parent shoot. Copyright 1999 Annalsof Botany Company Atlas cedar, extension growth, sylleptic branching, tree architecture, morphology.     

Quantitative analysis of the phenotypic variability of shoot architecture in two grapevine (Vitis vinifera) cultivars

BACKGROUND AND AIMS: Plant architecture and its interaction with agronomic practices and environmental constraints are determinants of the structure of the canopy, which is involved in carbon acquisition and fruit quality development. A framework for the quantitative analysis of grapevine (Vitis vinifera) shoot architecture, based on a set of topological and geometrical parameters, was developed for the identification of differences between cultivars and the origins of phenotypic variability. METHODS: Two commercial cultivars ('Grenache N', 'Syrah') with different shoot architectures were grown in pots, in well-irrigated conditions. Shoot topology was analysed, using a hidden semi-Markov chain and variable-order Markov chains to identify deviations from the normal pattern of succession of phytomer types (P0-P1-P2), together with kinematic analysis of shoot axis development. Shoot geometry was characterized by final internode and individual leaf area measurements. KEY RESULTS: Shoot architecture differed significantly between cultivars. Secondary leaf area and axis length were greater for 'Syrah'. Secondary leaf area distribution along the main axis also differed between cultivars, with secondary leaves preferentially located towards the basal part of the shoot in 'Syrah'. The main factors leading to differences in leaf area between the cultivars were: (a) slight differences in main shoot structure, with the supplementary P0 phytomer on the lower part of the shoot in 'Grenache N', which bears a short branch; and (b) an higher rate and duration of development of branches bearing by P1-P2 phytomers related to P0 ones at the bottom of the shoot in 'Syrah'. Differences in axis length were accounted for principally by differences in individual internode morphology, with 'Syrah' having significantly longer internodes. This trait, together with a smaller shoot diameter, may account for the characteristic 'droopy' habit of 'Syrah' shoots. CONCLUSIONS: This study highlights the architectural parameters involved in the phenotypic variability of shoot architecture in two grapevine cultivars. Differences in primary shoot structure and in branch development potential accounted for the main differences in leaf area distribution between the two cultivars. By contrast, shoot shape seemed to be controlled by differences in axis length due principally to differences in internode length.     

The effect of autumn N supply on the architecture of young peach (Prunus persica L.) trees

Irrigation and fertilisation were recently considered as useful tools to control tree shape, and reduce pruning costs. The role of the N reserves, which determined spring growth, was considered to be essential. We intended therefore to evaluate its effects on peach tree architecture. Four levels of N fertilisation were applied on 1-year-old trees, from the end of shoot growth to leaf fall. In subsequent spring, each bud fell into one of the ten classes of positions previously defined within the crown. Its development was followed weekly from burst to June. Fertilisation promoted growth until a threshold level, since no differences were evidenced between the three highest N treatments. Fall N did not affect burst but the further transformation of the buds into rosettes, proleptic or ramificated axes. Crown base was little affected. Fall N increased the number of proleptic axes on most median and upper positions. Axes lengthening and thickening were limited on the median positions, promoted at crown top. The variations concerned the mean internodes lengths, not the number of phytomers per axis. Sylleptic ramification was limited to the crown outer parts, and decreased with fall N. Treatment did neither affect the fruit dry weights, nor the ratio between the number of leaves and the number of fruits. Fruit number was proportioned to vegetative growth by blossoming and fruit set. We conclude that a moderate autumn fertilisation improved orchard productivity, but favoured vegetative growth in the crown outer parts. Additional pruning may therefore be required to control tree shape.     

Shaping the shoot: the relative contribution of cell number and cell shape to variations in internode length between parent and hybrid apple trees

Genetic control of plant size and shape is a promising perspective,particularly in fruit trees, in order to select desirable genotypes.A recent study on architectural traits in an apple progeny showedthat internode length was a highly heritable character. However,few studies have been devoted to internode cellular patterningin dicotyledonous stems, and the interplay between the two elementarycell processes that contribute to their length, i.e. cell divisionand elongation, is not fully understood. The present study aimedat unravelling their contributions in the genetic variationof internode length in a selection of F₁ and parent genotypesof apple tree, by exploring the number of cells and cell shapewithin mature internodes belonging to the main axes. The resultshighlighted that both the variables were homogeneous in samplescollected either along a sagital line or along the pith width,and suggest that cell lengthening was homogeneous during internodedevelopment. They allowed the total number of cells to be estimatedon the internode scale and opened up new perspectives for simplifyingtissue sampling procedures for further investigations. Differencesin internode length were observed between the genotypes, inparticular between the parents, and partly resulted from a compensationbetween cell number and cell length. However, genetic variationsin internode length primarily involved the number of cells,while cell length was more secondary. These results argue foran interplay between cellular and organismal control of internodeshape that may involve the rib meristem. Key words: Elongation, growth, histogenesis, Malusxdomestica Borkh, pith Received 2 January 2008; Revised 22 January 2008 Accepted 29 January 2008     

Comparison of Horizontal Spread of White Clover (Trifolium repensL.) Grown Under Two Artificial Light Sources Differing in their Content of Blue Light

The effects of blue light on the growth and development of differentbranches were studied to determine if variations in the bluelight supply modified the horizontal spread of white clover.Cuttings of white clover (‘Huia’) were grown for56 d in two controlled chambers under conditions similar exceptfor level of blue light (BL). The horizontal spread of whiteclover was characterised by the phyllochron, internode lengthand branching or flowering of different axes. The responsesto a reduction in BL were different according to axis degree:(1) on the main axis, lowering BL slightly decreased the phyllochronand had no effect on internode length or stolon length; (2)on older primary branches, it decreased their phyllochron, butreduced stolon length due to a reduction in internode length;(3) on secondary branches, it increased the phyllochron anddecreased internode length, triggering a strong reduction instolon length (up to 54%). Moreover, an increase in BL reversedthese effects. The switch to lower BL or higher BL had no effecton internode length of primary branches, but triggered changesin internode length of the main axis. These differential effectsof variations in BL on the morphogenesis of different branchesare discussed in relation to the clonal integration of variationsin light composition. Changes in the horizontal spread of whiteclover induced by BL could play a role, combined with responsesto low red:far-red ratio and low photosynthetic photon flux(PPF) in the strategy of horizontal colonisation and shade-avoidanceof white clover.Copyright 1998 Annals of Botany Company Blue light, branching, horizontal spread, morphogenesis, phyllochron, plant development, plant morphology, stolon growth,Trifolium repensL., white clover.     

Relationship between Sylleptic Branching and Components of Parent Shoot Development in the Peach Tree

Sylleptic branching of main axes was investigated in three peachtree cultivars ('Armking', 'Flavorcrest' and 'Silvergem') duringthe first year of growth. An axis was considered as made upof a series of metamers (internode, node, leaf and associatedbud) and its growth was divided into two components: the increaseof the number of metamers and the lengthening of the metamersthemselves (elongation). The relationship between branchingprobability, type of cultivar, growth components and date ofmetamer production was studied by logistic regression. Branchingprobability varied according to the cultivar, increased whenthe growth components did, and decreased if metamers appearedlate in the season. The logistic model fitted the data closelyand was validated on a data set that had not been used for estimatingthe parameters. Ninety-four percent of branched and 70% of unbranchedmetamers were correctly predicted by the logistic model. Forany given growth rate and date of metamer production, the mainaxes branched most and least often in the 'Flavorcrest' and'Silvergem' cultivars, respectively.Copyright 1994, 1999 AcademicPress Peach tree, Prunus persica (L.) Batsch, branching, syllepsis, shoot growth, quantitative analysis, logistic regression     

Limits to tree species invasion in pampean grassland and forest plant communities

Factors limiting tree invasion in the Inland Pampas of Argentina were studied by monitoring the establishment of four alien tree species in remnant grassland and cultivated forest stands. We tested whether disturbances facilitated tree seedling recruitment and survival once seeds of invaders were made available by hand sowing. Seed addition to grassland failed to produce seedlings of two study species, Ligustrum lucidum and Ulmus pumila, but did result in abundant recruitment of Gleditsia triacanthos and Prosopis caldenia. While emergence was sparse in intact grassland, seedling densities were significantly increased by canopy and soil disturbances. Longer-term surveys showed that only Gleditsia became successfully established in disturbed grassland. These results support the hypothesis that interference from herbaceous vegetation may play a significant role in slowing down tree invasion, whereas disturbances create microsites that can be exploited by invasive woody plants. Seed sowing in a Ligustrum forest promoted the emergence of all four study species in understorey and treefall gap conditions. Litter removal had species-specific effects on emergence and early seedling growth, but had little impact on survivorship. Seedlings emerging under the closed forest canopy died within a few months. In the treefall gap, recruits of Gleditsia and Prosopis survived the first year, but did not survive in the longer term after natural gap closure. The forest community thus appeared less susceptible to colonization by alien trees than the grassland. We conclude that tree invasion in this system is strongly limited by the availability of recruitment microsites and biotic interactions, as well as by dispersal from existing propagule sources.     

Correlative Growth in Young Terminalia superba in a Controlled Environment: Effect of the Leaves on Internode Elongation

Young Terminalia superba plants were cultivated in a controlledenvironment at the Phytotron. Effects of the excision of a youngleaf at definite elongation stages and at two given levels ofthe main axis were studied on the elongation of internodes inthese plants. Effects of the leaf did not seem to depend onits nodal position on the main axis but predominantly occurredon the immediate surrounding internodes. The excision of a youngleaf enhanced the growth rate of the internode located belowit and markedly decreased the elongation of the internode aboveit but slightly affected internode growth duration. This excisionenhanced the final length of the internode located below theleaf and decreased the final length of the internode locatedabove the removed leaf. Significant linear regressions werefound between the length of the excised leaf and the internodefinal lengths. Microscopic examination of epidermal cells ofcontrol and disrupted internodes revealed that the decreasedelongation after leaf excision could be attributed to reductionof cell divisions. The increased elongation after leaf excisioncould be attributed both to slight increase in the length ofcells (significant negative correlation was found with the lengthof the removed leaf) and to increase of cell divisions. Terminalia superba, leaf excision, cell division, internode elongation, correlative growth     

Morphological and histological differences in the development of dwarf mutants of sexual and somatic origin in diverse woody taxa

Morphological and histological observations were made on eight dwarf mutants arising either as seedlings from sexual reproduction or from somatic bud mutations (witches'-brooms in the crowns of normal trees). The most predominate morphological trait contributing to the expression of dwarfism in all taxa was the reduction of final internode lengths along the shoot axis. In taxa of sexual origin, with the exception of Prunus, there was a consistent reduction in the number of preformed leaves contained in the winter buds. In addition, in two taxa (Liquidambar and Tsuga) there was an almost complete absence of neoformed leaves and sylleptic branches on current year shoots. Conversely, in mutants of somatic origin there was no apparent reduction in the number of preformed leaves. Genetic dwarfness in this group resulted solely from decreases in final internode length. Significant differences in the cellular basis of dwarfism between mutants of different genetic origins are clearly evident. In dwarf trees arising from sexual reproduction, reduction in final internode length is attributed predominately to inhibition of mitotic activity in developing internodes resulting in highly significant decreases in final cell number, and not cell length. In mutants of somatic origin, the reduction in length of mature internodes results from a decrease in final cell length, rather than a decrease in cell number. Physiological mechanisms associated with the genetic expression of these morphogenetic differences are suggested.     

Gravimorphism in Trees: 1. Effects of Gravity on Growth and Apical Dominance in Fruit Trees

The effect of shoot-orientation on the vegetative growth andapical dominance of rootstocks of apply, cherry, plum, and blackcurrantwas investigated. In all the species studied the total annualextension growth of horizontally grown trees was markedly lessthan that of vertical trees, and the effect was accentuatedwhen the plants were rotated. In cherry and plum the reducedtotal growth was due both to a smaller number of internodesand also to reduced internode length; in apple and blackcurrantthe difference in extension growth was mainly due to differencesin internode length. In experiments with laternal shots trainedin various positions it was found that the apical dominancenormally exhibited by the uppermost shoot is only manifestedwhen it occupies a vertical position. The normal apical dominancerelationships can be reversed if the uppermost shoot is grownhorizontally and the second shoot vertically. Gravitationaleffects appear to play a role in the normal apical dominancerelationships of the tree. In all species there is a markedtendency for laterals to grow out on the upper side of horizontaland arched plants. The effects of training arched trees into various positionsare considered in relation to current concepts regarding therole of auxin in apical dominance and the redistribution ofauxin in horizontal organs; it is concluded that the outgrowthof laterals in arched shoots cannot be interpreted primarilyin terms of these current concepts. The observed results suggestthat the outgrowth of lateral buds occurs at the nearest pointto the roots at which the shoot is diverted from the verticalposition, and that nutrients are diverted to the highest upwardlydirected meristem. It is proposed that the term gravimorphismshould be applied to these diverse effects of gravity on plantgrowth.     

THE ROLE OF RESERVES IN LEAVES,BRANCHES, STEMS,AND ROOTS ON SHOOT GROWTH OF RED PINE

By defoliation, girdling, and various combinations of these before the growing season began, the role of foods in old needles, branches, main stems, and roots on shoot growth of 8-yr-old red pine (Pinus resinosa) trees was studied in northern Wisconsin. Defoliation and girdling, alone or in combination, reduced shoot growth but defoliation reduced shoot dry weight more than it reduced shoot elongation. Reductions in shoot growth due to treatment were in the following decreasing order: branch girdling + needle removal > girdling at stem base + needle removal > needle removal > branch girdling > base girdling. The old needles were the major source of food for shoot growth and accounted for four-fifths or more of all shoot growth. The combined reserves in the branches, main stem, and roots accounted for less than 15% of shoot growth. The contribution of reserves from tissues other than old leaves was in the following order: branches > main stem > roots. Defoliation weakened apical dominance relations. In defoliated trees, many secondary axes elongated more than the terminal leader, and secondary axes in lower whorls often grew more than those in upper ones. The data suggest an important role of nutritional factors in correlative growth inhibition.     

The role of the leaves in the regulation of internode elongation in Phaseolus vulgaris

Time-course patterns of leaf and internode elongation were studied in bean plants. Each leaf started its main elongation period when the leaf below reached half of its final length. The onset of leaf unfolding was nearly synchronous with the initiation of the elongation of the subjacent internode. Excision of young leaves increased the rate of stem elongation as a result of an earlier unfolding of the next upper leaves and the concomitant advancement in the elongation of their subjacent internodes. IAA or NAA (1% in lanolin) suppressed the enhancement effects of leaf excision on leaf and internode elongation. The excision of a young leaf increased the final length of internodes located below it, and at the same time decreased the final length of the internodes located above the excised leaf. The reduction was greater the younger the internode. Differences in internode elongation after leaf excision were related to changes during internode ontogenesis in their relative response to the availability of assimilates on the one hand, and on the other hand to hormonal factors transported acropetally from the young leaves to the growing internodes.     

Spatial variability in oviposition damage by periodical cicadas in a fragmented landscape

Effects of the periodical cicada (Magicicada spp.) on forest dynamics are poorly documented. A 1998 emergence of M. cassini in eastern Kansas led to colonization of a fragmented experimental landscape undergoing secondary succession. We hypothesized that per-tree rates of oviposition damage by cicadas would reflect: (1) distance from the source of the emergence, (2) patch size, and (3) local tree density. Ovipositing females displayed clear preferences for host species and damage incidence showed predictable spatial patterns. Two species (smooth sumac, Rhus glabra, and eastern red cedar, Juniperus virginiana) were rarely attacked, whereas others (rough-leaved dogwood, Cornus drummondii; slippery elm, Ulmus rubra; box elder, Acer negundo, and honey locust, Gleditsia triacanthos) were strongly attacked. The dominant early successional tree, dogwood, received on average the most attacks. As predicted, attacks per stem declined strongly with distance from the emergence source, and with local stem density (a "dilution" effect). Contrary to expectations, there were more attacks per stem on larger patches. Because ovipositing cicadas cut damaging slits in host tree branches, potentially affecting tree growth rate, competitive ability, and capacity to reproduce, cicada damage could potentially influence spatial variation in secondary succession.     

Co-ordinated growth between aerial and root systems in young apple plants issued from in vitro culture

BACKGROUND AND AIMS: In several species exhibiting a rhythmic aerial growth, the existence of an alternation between root and shoot growth has been demonstrated. The present study aims to investigate the respective involvement of the emergence of new organs and their elongation in relation to this phenomenon and its possible genotypic variation in young apple plants. METHODS: Two apple varieties, X6407 (recently named 'Ariane') and X3305 ('Chantecler' x 'Baujade'), were compared. Five plants per variety, issued from in vitro culture, were observed in minirhizotrons over 4 months. For each plant, root emergence and growth were observed twice per week. Growth rates were calculated for all roots with more than two segments and the branching density was calculated on primary roots. On the aerial part, the number of leaves, leaf area and total shoot length were observed weekly. KEY RESULTS: No significant difference was observed between varieties in any of the final characteristics of aerial growth. Increase in leaf area and shoot length exhibited a 3-week rhythm in X3305 while a weaker signal was observed in Ariane. The primary root growth rate was homogeneous between the plants and likewise between the varieties, while their branching density differed significantly. Secondary roots emerged rhythmically, with a 3-week and a 2-week rhythm, respectively, in X3305 and 'Ariane'. Despite a high intra-variety variability, significant differences were observed between varieties in the secondary root life span and mean length. A synchronism between leaf emergence and primary root growth was highlighted in both varieties, while an opposition phase was observed between leaf area increments and secondary root emergence in X3305 only. CONCLUSION: A biological model of dynamics that summarizes the interactions between processes and includes the assumption of a feedback effect of lateral root emergence on leaf emergence is proposed.     

Sylleptic branching in winter-headed apple (<Emphasis Type="Italic">Malus × domestica)</Emphasis> trees: accession-dependent responses and their relationships with other tree architectural characteristics

Well-feathered apple trees are essential for commercial orchards to optimize yields. However, most cultivars do not form these sylleptic branches readily in commercial nurseries due to high apical dominance. Several treatments exist to promote their formation in the nurseries, one of which is heading. However, not all cultivars are expected to react similarly to these treatments. We studied the branching response of 155 genotypes following heading and its relation to other architectural traits as a function of the cultivar’s genetic background. Trees were grown for two consecutive years after grafting in a nursery, and the main axes were headed in the winter following growth in year 1. After heading, a single shoot was retained when growth resumed in the second year. Plant architectural traits such as growth of the main shoot, internode length, sylleptic branching, etc. were measured before and after heading and were statistically compared using a mixed model. This model showed the effect of heading for all architectural traits studied. In addition significant genotype and genotype × treatment interactions were found. In general, genotypes that showed a more vigorous growth during the first year also reacted more vigorously to heading. Accounting for the genetic substructure of this collection, no clear distinction in tree response could be found except in a small group of individuals that belonged to an F1 mapping population. This study shows that heading is not favorable for all genotypes to promote sufficient sylleptic branching and that other methods are needed to promote branching in these accessions.     

Nutrient partitioning and storage in arid-zone forests of the mangroves Rhizophora <Emphasis Type="Italic">stylosa</Emphasis> and <Emphasis Type="Italic">Avicennia</Emphasis><Emphasis Type="Italic">marina</Emphasis>

Mangrove partitioning and storage of macronutrients and trace metals were examined in different arid coastal settings of Western Australia. Total living biomass in three Rhizophora stylosa forests, which ranged from 233 to 289 t DW ha^-1, was significantly greater than biomass in three Avicennia marina forests (range: 79-155 t DW ha^-1). Although prop roots and stems were the largest single tree components for R. stylosa and A. marina, respectively, most nutrients were stored in leaves and living roots of both species. However, only a small fraction of the total nutrient pool was vested in tree biomass; the vast bulk was in soils. A large below-ground pool of dead fine roots was identified at all stands, equivalent to 36-88% DW of total living tree biomass. The amount of Ca, S, Cl, Na, Si, Fe, Mn, Zn, B, Mo and Cu vested in dead roots of both species was greater than in the total living tree biomass. The proportion of Fe and S vested in live and dead roots was exceptionally large, consistent with previous evidence of metal plaques on mangrove roots. Sulphur, iron and zinc in dead roots of both species constituted the bulk of these metals. R. stylosa trees preferentially accumulated more Mg, S, Cl, Na, Si, Fe, Mn, B and Mo than A. marina trees. Proportionally greater storage of P, N, Ca, K, Cu and Zn occurred in two of the three A. marina forests. Foliar concentrations of Mg, S, Mn, B and Mo in mangrove leaves were at the high end of the range reported for other tropical trees, but other elemental concentrations were at the low or mid-range. Nitrogen limitation in these forests is implied by a positive correlation between total tree N and net canopy production and by a lower percentage of ecosystem N in tree biomass as compared with other forests. Unlike terrestrial forests where a large proportion of nutrient capital is vested in floor litter, most elements in these mangrove forests are stored in dead roots. A large reservoir of dead roots below the forest floor may serve as a conservation mechanism, particularly in such arid oligotrophic environments.     

Growth and Nutrient Retranslocation in Needles of Radiata Pine in Relation to Nitrogen Supply

The effects of N application on tree growth and the retranslocationof N, P, and K from young needles to new growth were examinedin young radiata pine (Pinus radiata D. Don) trees. Nitrogen fertilization increased the number and size of needles,rates of shoot production, stem volume growth and tree biomass.Foliar N and P contents (µg per needle) fluctuated ina cyclic fashion with prominent phases of accumulation, retranslocationand replenishment. The patterns of these fluctuations in controland N-fertilized trees were similar, although the fluxes ofN, P and K in and out of needles were increased by N fertilization.Greater translocation (g per tree) of N and K from needles ofN fertilized trees occurred because fertilization increasedthe needle weight and the proportion of N and K retranslocatedfrom individual needles. Nitrogen fertilization increased theretranslocation of P largely as a result of higher needle mass.Trees supplied with more than adequate amounts of P in the soilretranslocated up to 58 per cent of the initial pool of P fromyoung needles. The periods of high retranslocation coincidedwith periods of high concentrations of soil mineral N and withshoot production. Conversely, the periods of rapid replenishmentof N and P into the needles coincided with the time of slowshoot growth and low concentration of soil mineral N. The growthrate of trees, rather than the availability of nutrients inthe soil was the main factor controlling retranslocation. For radiata pine, retranslocation from needles is not a mechanismspecific for coping with low soil fertility. It seems to bea mechanism which enhances the nutrient supply to apical growingpoints, especially during periods of flushing. Pinus radiata, nitrogen supply, shoot growth, nutrient fluctuations and retranslocation, nutrient use and adaptation