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PubMed2004

A Pinus radiata AAA-ATPase, the expression of which increases with tree ageing.

Age-dependent changes in gene expression profiles were studied in vegetative Pinus radiata buds by means of differential display. Among several candidate cDNAs, a 327 bp fragment that shows high homology with an Arabidopsis thaliana 20S proteasome ATPase designated RPT5a was found. Northern hybridization confirmed that the accumulation of this transcript increases with tree ageing, suggesting a possible role of this AAA-ATPase gene in development-related specific proteolysis.

Adenosine Triphosphatases↗

PubMed2005

Phototropic response induced by wind loading in Maritime pine seedlings (Pinus pinaster Aït.).

Both woody and herbaceous plant species are known to respond to wind loading, with consequences for growth and morphology. Wind has usually been classified as a mechanical stress which is detrimental to plant growth. Few experiments exist whereby plants and, in particular, woody species are exposed to wind, as opposed to mechanical perturbation by touching, flexing or shaking. Such experiments have always been short term and often carried out in wind tunnels in a controlled greenhouse environment. This study introduces an experiment to test the responses of Maritime pine (Pinus pinaster Aït.) seedlings to recurrent and short wind loading in the field, over two growing seasons. These experiments provide evidence that periodic short-term exposure to wind can induce phototropic responses in the early stage of pine seedlings' development. An interpretation is proposed in terms of efficiency to light tracking and hypotheses are discussed concerning the underlying physiological process.

Phototropism↗

PubMed2005

A mechanical analysis of the relationship between free oscillations of Pinus pinaster Ait. saplings and their aerial architecture.

The aim of this study was to investigate the influence of aerial architecture on the dynamic characteristics of young maritime pines (Pinus pinaster Ait.) using a mechanistic approach. For this purpose, three 4-year-old saplings with prominent differences in their branching patterns were submitted to free oscillation tests. The tests were carried out with different methods and directions of mechanical loading in order to initiate the movement of each sapling. The oscillations of the different architectural elements, i.e. stem and branches of different topological order, were measured with inclinometers and strain gauges fixed to saplings. Successive pruning of the architectural elements was carried out to evaluate their relative influence on the dynamic characteristics of the trees. The aerial systems were digitized before the mechanical tests in order to use 3D visualization techniques and to make architectural analyses of the crown structure. Two distinct modes of deformation were detected during free oscillations. The natural swaying frequency ranged from 0.6-0.8 Hz for the saplings tested at the same period of the year. The frequency variations were partly explained by the morphological differences of the experimental subjects. The motions of the axes were found to depend on their topology, i.e. the movement of the axes of a given branching order was forced by the movement of their respective bearing axis. The axes of third branching order had a significant and negative effect on the damping of the natural deformation mode. Results point out the major role played by foliage, qualitatively and quantitatively, on the damping of tree motions and on coupling the motions of the crown components.

Biomechanical Phenomena↗

PubMed2006

Uptake, metabolism and distribution of organic and inorganic nitrogen sources by Pinus sylvestris.

Although an increasing number of studies show that many plant species have the capacity to take up amino acids from exogenous sources, the importance of such uptake for plant nitrogen nutrition is largely unknown. Moreover, little is known regarding metabolism and distribution of amino acid-N following uptake or of the regulation of these processes in response to plant nitrogen status. Here results are presented from a study following uptake, metabolism, and distribution of nitrogen from NO(3)(-) NH(4)(+), Glu, or Ala in Scots pine (Pinus sylvestris L). In a parallel experiment, Ala uptake, processing, and shoot allocation were also monitored following a range of pretreatments intended to alter plant C- and N-status. Uptake data, metabolite profiles, N fluxes through metabolite pools and tissues, as well as alanine aminotransferase activity are presented. The results show that uptake of the organic N sources was equal to or larger than NH(4)(+) uptake, while NO(3)(-) uptake was comparatively low. Down-regulation of Ala uptake in response to pretreatments with NH(4)NO(3) or methionine sulphoximine (MSX) indicates similarities between amino acid and inorganic N uptake regulation. N derived from amino acid uptake exhibited a rapid flux through the amino acid pool following uptake. Relative shoot allocation of amino acid-N was equal to that of NH(4)(+) but smaller than for NO(3)(-) Increased N status as well as MSX treatment significantly increased relative shoot allocation of Ala-N suggesting that NH(4)(+) may have a role in the regulation of shoot allocation of amino acid-N.

Alanine↗

PubMed2006

Genetic structure and evolutionary history of a diploid hybrid pine Pinus densata inferred from the nucleotide variation at seven gene loci.

Although homoploid hybridization is increasingly recognized as an important phenomenon in plant evolution, its evolutionary genetic mechanisms are poorly documented and understood. Pinus densata, a pine native to the Tibetan Plateau, represents a good example of a homoploid hybrid speciation facilitated by adaptation to extreme environment and ecological isolation from the parents. Its ecologically and reproductively stabilized nature offers excellent opportunity for studying genetic processes associated with hybrid speciation. In this study, we investigated the levels and patterns of nucleotide variation in P. densata and its putative parents. Haplotype composition, gene genealogies, and the levels and patterns of nucleotide variation gave further support to the hybrid nature of P. densata. Allelic history, as revealed by our data, suggests the ancient nature of the hybrid preceding elevation of the Tibetan Plateau. We detected more deviations from neutrality in P. densata than in the parental species. Thus, at least some of the evolutionary forces that have shaped the genetic variation in P. densata are likely to be different from those acting upon parental species. We speculate that when populations of P. densata invaded new territories, they had elevated rates of response to selection in order to develop traits that help them to survive and adapt in the new environments.

Diploidy↗

PubMed1998

Loblolly pine (Pinus taeda L.) contains multiple expressed genes encoding light-dependent NADPH:protochlorophyllide oxidoreductase (POR).

NADPH:protochlorophyllide oxidoreductase (POR) catalyzes the light-dependent reduction of protochlorophyllide (Pchlide) to chlorophyllide (Chlide), a key regulatory step in chlorophyll biosynthesis. In most angiosperms, POR is encoded by a small nuclear gene family, containing at least two differentially-expressed genes designated porA and porB. We have demonstrated that the PORs of loblolly pine (Pinus taeda L.), a gymnosperm, are encoded by a large multigene family, composed of two distinct subfamilies encoding porA and porB genes similar to those previously described in angiosperms. DNA gel blot analysis of genomic DNA showed that the two por subfamilies of loblolly pine have duplicated at different rates, with the porA subfamily containing two members, and the porB subfamily containing at least 11 potential members. DNA sequence analysis and gel blot hybridization studies also showed that a subset of the por genes present in the loblolly pine genome are pseudogenes. Based on the results of 5'- and 3'-RACE analysis, it appears that multiple porA and porB genes are expressed in loblolly pine cotyledons and stems during development. Using gene specific probes, no difference was observed in the steady-state levels of the different porA and porB transcripts in cotyledons of dark-grown seedlings before and following illumination. However, the steady state levels of the porA and porB transcripts were found to increase at different rates in the stems of dark-grown seedlings following illumination. The phylogenetic relationship between the por gene family members in P. taeda and other pine species and the potential significance of the two por subfamilies to the evolution of por gene function are discussed.

Amino Acid Sequence↗

PubMed2005

Casparian strips in needles are more solute permeable than endodermal transport barriers in roots of Pinus bungeana.

The structure and development of endodermal Casparian strips in Pinus bungeana needles and roots were studied by scanning electron microscopy and fluorescence microscopy. Primary pit fields (PFs) frequently occurred in radial walls of Casparian strips isolated from needles, whereas PFs were never detected in Casparian strips from roots. Formation of Casparian strips in needles as well as roots started at the outer parts of the radial walls and they finally occupied the entire radial walls of the endodermis. Fourier transform infrared (FTIR) spectroscopy of Casparian strips isolated from roots revealed significant absorption bands characteristic for suberin. However, in Casparian strips of needles, evidence for suberin was rarely detected by FTIR spectroscopy. The apoplastic permeability of Casparian strips in needles and roots was probed by the apoplastic tracers calcofluor and berberine. Casparian strips in roots efficiently blocked the apoplastic transport (AT) of calcofluor and berberine. Casparian strips in needles blocked the AT of calcofluor, but diffusion of berberine was not inhibited and berberine thiocyanate crystals were detectable in the vascular tissue of the needles. From the data presented, it must be concluded that Casparian strips in needles, which are characterized by the absence of suberin, are more solute permeable compared with Casparian strips in roots.

Biological Transport↗

PubMed2006

The non-host pathogen Botrytis cinerea enhances glucose transport in Pinus pinaster suspension-cultured cells.

Botrytis cinerea is the causal agent of grey mould disease and a non-host necrotrophic pathogen of maritime pine (Pinus pinaster). Recent evidence suggests that pathogen challenge can alter carbon uptake in plant cells; however, little is known on how elicitor-derived signalling pathways control sugar transport activity. P. pinaster suspended cells are able to absorb D-[14C]glucose with high affinity, have an H+-dependent transport system (Km, 0.07 mM; Vmax, 1.5 nmol min(-1) mg(-1) DW), are specific for D-glucose, D-fructose, D-galactose and D-xylose, and are subject to glucose repression. When elicited by B. cinera spores, suspended cells exhibit calcium-dependent biphasic reactive oxygen species (ROS) production, the second burst also being dependent on NADPH oxidase, mitogen-activated protein kinase (MAPK), and de novo transcription and protein synthesis. Challenging suspended cells incubated in sugar-free medium resulted in an up to 3-fold increase in glucose transport capacity over non-elicited cultures 24 h after elicitation, and a 14-fold increase over elicited cells incubated with 2% glucose. Enhanced glucose uptake depended on NADPH oxidase and calcium influx, but not MAPK. In contrast, the increase of glucose transport activity induced by sugar starvation was dependent on the activation of MAPK but not NADPH oxidase. Both responses appeared to be dependent on de novo transcription and protein synthesis.

Biological Transport↗

PubMed2000

A mycobacterium isolated from tissue cultures of mature Pinus sylvestris interferes with growth of Scots pine seedlings.

We isolated a rapidly growing, pigment-producing mycobacterium from senescent tissue cultures derived from mature Scots pine (Pinus sylvestris L.). The bacterium was found in three senescent suspension cultures and in a senescent protoplast culture. Growth of Scots pine cells had ceased in all of these cultures. Exogenous contamination was eliminated by rigorous surface sterilization of the buds with hypochlorite before aseptic removal of the bud scales. Based on biochemical and physiological properties and DNA sequence comparisons, the isolated mycobacterium did not belong to any known species. Its sequence most closely resembled those of Mycobacterium obuense (97%) and M. aichiense (96%). Tissue browning was frequently observed in callus or suspension culture of Scots pine. Because the effect of the mycobacterium on growth of undifferentiated tissues that were browning was difficult to evaluate, we applied the bacterium to Scots pine seeds in aseptic conditions. Seedlings grown in the presence of the mycobacterium had shorter hypocotyls than control seedlings and seedlings cocultivated with a Pseudomonas strain known to be harmless to plants. However, hypocotyl growth of seedlings cocultivated with another mycobacterium, M. chlorophenolicum, was similar to that observed in the presence of the isolated mycobacterium. Phenylalanine ammonia-lyase activity of seedlings cocultivated with the mycobacterium isolate was significantly higher than that of control seedlings or seedlings cocultivated with M. chlorophenolicum or Pseudomotnas fluorescens. We believe that this is the first report of the isolation of mycobacteria from tissue cultures of a tree. Our finding that the mycobacterium may interfere with the growth of Scots pine in vitro warrants further study.

Culture Techniques↗

PubMed2001

Nitrogen forms in bark, wood and foliage of nitrogen-fertilized Pinus sylvestris.

Cycling of soluble non-protein N compounds is thought to be indicative of the N-nutritional status of trees. We determined the major N forms in bark, wood and foliage and estimated the dependence of prevalent N forms on N availability in Pinus sylvestris L. trees from northern Sweden. Trees subjected to severe N limitation and trees that had been fertilized with an average 64 kg N ha(-1) year(-1) for 25 years were analyzed. Bark and wood samples were collected by tangentially cryo-sectioning the trunk into 30-microm thick sections, from the bark to the functional xylem. Soluble amino compounds were extracted from the sections for analysis. Sap samples from twigs were obtained by centrifugation, and bark samples from twigs were obtained by tissue extraction. In both needles and bark, arginine dominated the amino-N pool. Because arginine concentrations in needles increased with N fertilization, arginine dominance of the amino-N pool in needles was higher in N-fertilized trees than in control trees. In bark, N fertilization resulted in a large increase in glutamine concentration, so that glutamine accounted for a larger proportion of the amino-N pool in bark in N-fertilized trees than in control trees. Glutamine dominated the amino-N pool in wood of control trees. Nitrogen fertilization resulted in an increased proportion of arginine in the wood amino-N pool. We conclude that the composition of the amino-N pools in bark, wood and foliage is highly sensitive to N supply. The composition of the amino-N pools can contribute to the regulation of tree N-nutritional status, which is mediated by shoot to root signalling by long-distance transport of amino compounds.

Cycadopsida↗

PubMed2001

Inducible expression of the heterologous PAL2 promoter from bean in white pine (Pinus strobus) transgenic cells.

To elucidate heterologous promoter function in gymnosperms, we introduced the bean phenylalanine ammonia-lyase-beta-glucuronidase (PAL2-GUS) gene fusion into white pine (Pinus strobus L.). Over 15 lines were produced and integration of Agrobacterium T-DNA was confirmed by Southern analysis. Induction of the reporter gene was detected in all of the lines tested following UV illumination. In contrast, a weak but constant induction was seen in only a few lines following treatment with salicylic acid (SA) or jasmonic acid (JA). However, pretreatment of suspension cultures with SA or JA enhanced the induction of PAL2-GUS expression by UV irradiation. This specific enhancement or potentiation was reduced by 50% by treating the cells with indomethacin, an inhibitor of phospholipase activity, suggesting that the observed potentiation of UV induction involves the octadecanoid pathway. The UV induction was completely abolished by treating the cells with okadaic acid, an inhibitor of phosphatase activity. Thus, the induction of the heterologous PAL2 promoter from bean is consistent with the induction of phenylalanine ammonia-lyase (PAL) in angiosperms. Furthermore, our findings suggest that conifers, although phylogenetically distant to angiosperms, share some conserved promoter elements and some signal transduction mechanisms for UV-light perception.

Artificial Gene Fusion↗

PubMed2001

Seasonal and interannual variations in carbon isotope discrimination in a maritime pine (Pinus pinaster) stand assessed from the isotopic composition of cellulose in annual rings.

Stable carbon isotope composition (delta; per thousand) was measured on cellulose extracted from maritime pine (Pinus pinaster Aït.) tree rings to investigate inter-tree and interannual variability (7 trees, 20 rings per tree, each ring divided into early and late wood). A model of stand primary production coupled to water balance was used to calculate the stand annual water-use efficiency (WUE). Inter-tree variability in discrimination (maximum 2.88 per thousand in late wood in 1989, 2.69 per thousand in early wood in 1983) was as large as interannual variation (maximum 2.72 per thousand in late wood, 2.05 per thousand in early wood). Tree size did not explain these differences. Relationships were found between annual discrimination and climate variables such as annual rainfall, summer temperature and vapor pressure deficit (VPD). Higher correlations were found with late wood discrimination. Early wood discrimination was shown to be related to previous-year late wood discrimination. Late wood discrimination was also related to soil water availability. Stand annual WUE was only weakly related to tree ring carbon discrimination.

Carbon Isotopes↗

PubMed2001

Canopy position and needle age affect photosynthetic response in field-grown Pinus radiata after five years of exposure to elevated carbon dioxide partial pressure.

Photosynthesis of tree seedlings is generally enhanced during short-term exposure to elevated atmospheric CO2 partial pressure, but longer-term studies often indicate some degree of photosynthetic adjustment. We present physiological and biochemical evidence to explain observed long-term photosynthetic responses to elevated CO2 partial pressure as influenced by needle age and canopy position. We grew Pinus radiata D. Don. trees in open-top chambers for 5 years in sandy soil at ambient (36 Pa) and elevated (65 Pa) CO2 partial pressures. The trees were well watered and exposed to natural light and ambient temperature. In the fourth year of CO2 exposure (fall 1997), when foliage growth had ceased for the year, photosynthetic down-regulation was observed in 1-year-old needles, but not in current-year needles, suggesting a reduction in carbohydrate sink strength as a result of increasing needle age (Turnbull et al. 1998). In 5-year-old trees (spring 1997), when foliage expansion was occurring, photosynthetic down-regulation was not observed, reflecting significantly large sinks for carbohydrates throughout the tree. Net photosynthesis was stimulated by 79% in trees growing in elevated CO2 partial pressure, but there was no significant effect on photosynthetic capacity or Rubisco activity and concentration. Current-year needles were more responsive to elevated CO2 partial pressure than 1-year-old needles, exhibiting larger relative increases in net photosynthesis to elevated CO2 partial pressure (98 versus 64%). Lower canopy and upper canopy leaves exhibited similar relative responses to growth in elevated CO2 partial pressure. However, needles in the upper canopy exhibited higher net photosynthesis, photosynthetic capacity, and Rubisco activity and concentration than needles in the lower canopy. Given that the ratio of mature to juvenile foliage mass in the canopy will increase as trees mature, we suggest that trees may become less responsive to elevated CO2 partial pressure with increasing age. We conclude that tree response to elevated CO2 partial pressure is based primarily on sink strength and not on the duration of exposure.

Carbon Dioxide↗

PubMed2001

Multivariate patterns of biochemical responses of Pinus ponderosa trees at field plots in the San Bernardino Mountains, southern California.

Most environmental stress conditions promote the production of potentially toxic active oxygen species in plant cells. Plants respond with changes in their antioxidant and photoprotective systems. Antioxidants and pigments have been widely used to measure these responses. Because trees are exposed to multiple man-made and natural stresses, their responses are not reflected by changes in single stress markers, but by complex biochemical changes. To evaluate such response patterns, explorative multivariate statistics have been used. In the present study, 12 biochemical variables (chloroplast pigments, state of the xanthophyll cycle, alpha-tocopherol, ascorbate and dehydroascorbate, glutathione and oxidized glutathione) were measured in previous-year needles of field-grown Pinus ponderosa Dougl. ex Laws. The trees were sampled in two consecutive years in the San Bernardino Mountains in southern California, where a pollution gradient is overlaid by gradients in natural stresses (drought, altitude). To explore irradiance effects, needle samples were taken directly in the field (sun exposed) and from detached, dark-adapted branches. A principal component analysis on this data set (n = 80) resulted in four components (Components 1-4) that explained 67% of the variance in the original data. Component 1 was positively loaded by concentrations of alpha-tocopherol, total ascorbate and xanthophyll cycle pools, as well as by the proportion of de-epoxides in the xanthophyll cycle. It was negatively loaded by the proportion of dehydroascorbate in the ascorbate pool. Component 2 was negatively loaded by chlorophyll concentrations, and positively loaded by the ratios of lutein and beta-carotene to chlorophyll and by the de-epoxidation state of the xanthophyll cycle. Component 3 was negatively loaded by GSH concentrations and positively loaded by the proportions of GSSG and tocopherol concentrations. Component 4 was positively loaded by neoxanthin and negatively loaded by beta-carotene. The four components could be assigned to the concerted action of the biochemical protection system: high scores on Component 1 represent highly activated antioxidative defense, changes in pigment composition are represented in Components 2 and 4, and the glutathione system, which is important for antioxidant regeneration, is represented in Component 2. Although Component 1 scores were generally higher (indicating activation of antioxidant defense) in light-adapted needles relative to dark-adapted needles, they were also site dependent with increased scores at sites with less pollution, but higher natural stress impacts. High scores of Components 2 and 3 at the highest elevation site, which was only moderately polluted, indicated an increase in photoprotection by pigments and activation of the glutathione system. Significant differences between light- and dark-adapted needles in Components 2 and 3 were only found at the site with the highest pollution. Use of accumulated variables (components) instead of single biochemical variables enabled recognition of response patterns at particular sites and a better comparison with results of other studies is expected. Typical response patterns could be assigned to particular environmental stress combinations, providing a means of assessing potential biological risks within individual forest stands.

Air Pollution↗

PubMed2001

Fungal auxin overproduction affects the anatomy of Hebeloma cylindrosporum-Pinus pinaster ectomycorrhizas.

We studied the effect of fungal auxin overproduction on the growth polarity of cortical cells in pine mycorrhizas by comparing the anatomy of Pinus pinaster (Ait.) Sol. mycorrhizas formed by an IAA-overproducing mutant of Hebeloma cylindrosporum Romagnesi or by the corresponding wild type with non-mycorrhizal short roots. Both wild- type and mutant strains induced an increase in root diameter that was mostly a result of the influence of the fungus on root cortical development. Both strains affected growth polarity of P. pinaster cortical cells and induced a change in their shape. The main modifications were a large reduction in axial diameter and an increase in the radial diameter of the cortical cells. The modifications were more marked with the mutant than with the wild type. The mutant induced a 43% reduction in cortical cell elongation and a 35% increase in radial diameter, whereas the corresponding changes induced by the wild type were 30 and 10%, respectively. The volume of cortical cells in mature mycorrhizas was generally lower than in uninoculated short roots indicating that wild-type and mutant strains induced a reorientation of cortical cell growth but did not induce an increase in turgor pressure of the cells. Immunolocalization allowed visualization of alpha-tubulin in root cortical cells, but no obvious modification in alpha-tubulin distribution was detected as a consequence of symbiosis establishment. Likewise, cytochemical localization of polysaccharides in cortical cell walls did not show significant modification following symbiosis establishment and Hartig net formation. The only noticeable modification was a reduction in cortical cell wall thickness in mycorrhizas compared with uninoculated short roots. The possible involvement of fungal auxin in the observed modifications is discussed.

Basidiomycota↗

PubMed2002

Phosphorus affects growth and partitioning of nitrogen to Rubisco in Pinus pinaster.

We tested the hypothesis that photosynthetic and growth responses to phosphorus (P) are functions of differences in the partitioning of nitrogen (N) among different compounds, particularly ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco). We tested this hypothesis in: (a) a greenhouse experiment with mycorrhizal seedlings of Pinus pinaster Ait. grown in sand culture for 4 months with six factorial combinations of N (0.125 and 2.0 mM) and P (0.02, 0.08 and 0.34 mM); and (b) a field study in which P was applied at five rates (up to 175 kg ha-1) to 2-year-old P. pinaster growing on P-deficient siliceous sand. After 4 months of nutrient addition or 2 years after fertilizer application, we measured light-saturated rates of photosynthesis, growth, and N and P allocation in needles. Growth of P. pinaster increased significantly with increasing concentrations of P, as did the concentration of P in needles. Concentrations of P and Rubisco were positively related, whereas those of N and Rubisco were unrelated. At low-P supply, the Rubisco/Chl ratio varied between 8.5 and 12 mmol mol-1. With P supply in excess of requirement (needle N:P ratio = 2-12) the Rubisco/Chl ratio increased to between 24 and 26 mmol mol-1. Rates of light-saturated photosynthesis were unaffected by P supply because adequate concentrations of P were maintained in plants in all treatments. Orthophosphate accumulated in needles of plants receiving a high P supply, which may allow growth to continue for periods under P deficiency, provided that other nutrients also accumulate. In the case of N, Rubisco may fill this role.

Amino Acids↗

PubMed2002

Response of a mature Pinus laricio plantation to a three-year restriction of water supply: structural and functional acclimation to drought.

The response of mature forest stands to a reduction in water availability has received little attention. In particular, the extent to which a short-term reduction in gas exchange can be alleviated in the long-term by acclimation processes is not well understood. We studied the impact of a severe reduction in water availability on the water relations and growth of 35-year-old Pinus laricio Poiret. trees in a replicated experiment. Sapwood and needle increments, soil and tree water status, stand transpiration, xylem embolism and plant hydraulic architecture were monitored over a 3-year period in control and drought-treated plots. Needle length was reduced in drought-treated trees by 30, 19 and 29%, and sapwood increments by 50, 27 and 24% over the 3 years. Drought did not result in tree mortality or in extensive xylem embolism or foliage dieback. On the contrary, a conservative water-use strategy was observed, because minimum leaf water potentials did not differ between treatments or over the season. Plant hydraulic resistance was also unaffected by restricted water availability. Stand transpiration was strongly reduced by drought treatment over the summer, but not during the winter, despite significant differences in leaf area between control and drought-treated trees, implying higher transpiration rates per unit leaf area in the droughted plants. This suggests that water transport capacity, rather than the amount of leaf area, controlled stand transpiration, which is at variance with expectations based on experiments with seedlings and short-term experiments with mature trees.

Disasters↗

PubMed2002

Dependence of needle architecture and chemical composition on canopy light availability in three North American Pinus species with contrasting needle length.

Morphology and chemical composition of needles of shade-intolerant southern conifers (Pinus palustris Mill. (mean needle length +/- SD = 29.1 +/- 4.1 cm), P. taeda L. (12.3 +/- 2.9 cm) and P. virginiana Mill. (5.1 +/- 0.8 cm)) were studied to test the hypothesis that foliage acclimation potential to canopy light gradients is generally low for shade-intolerant species, and in particular, because of mechanical limitations, in species with longer needles. Plasticity for each needle variable was defined as the slope of the foliar characteristic versus irradiance relationship. A novel geometrical model for needle area and volume calculation was employed for the three-needled species P. palustris and P. taeda. Needle thickness (T) strongly increased, but width (W) was less variable with increasing daily integrated quantum flux density averaged over the season (Q(int)), resulting in changes in cross-sectional needle shape that were manifested in a positive relationship between the total to projected needle area ratio (A(T)/A(P)) and Q(int) in the three-needled species. In contrast, cross-sectional needle geometry was only slightly modified by irradiance in the two-needled conifer P. virginiana. Needle dry mass per unit total needle area (M(T)) was positively related to Q(int) in all species, leading to greater foliar nitrogen contents per unit area at higher irradiances. Separate examination of the components of M(T) (density (D) and the volume (V) to A(T) ratio; M(T) = DV/A(T)) indicated that the positive effect of light on M(T) resulted solely from increases in V/A(T), i.e., from increases in the thickness of foliage elements. Foliar chlorophyll content per unit mass increased with increasing Q(int), allowing an improvement in light-harvesting efficiency in low light. The variables characterizing needle material properties (D, the dry to fresh mass ratio, and needle carbon content per unit mass) were generally independent of Q(int), suggesting that needles were less stiff and had greater tip deflections under their own weight at lower irradiances because of smaller W and T. Comparisons with the literature revealed that plasticity in foliar characteristics tended to be lower in the studied shade- intolerant species than in shade-tolerant conifers, but plasticity among the investigated species was unaffected by needle length. However, we argue that, because of mechanical limitations, plastic changes in needle cross section in response to low irradiance may decrease rather than increase light-interception efficiency in long-needled species.

Biomass↗