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SSR markers for Quercus suber tree identification and embryo analysis.

Three Quercus simple sequence repeat (SSR) markers were amplified by polymerase chain reaction (PCR) from nuclear DNA extracts of trees and in vitro-induced haploid embryos from anther cultures of Quercus suber L. These markers were sufficiently polymorphic to identify 10 of 12 trees located in two Spanish natural areas. The same loci have been analyzed in anther-derived haploid embryos showing the parental tree allele segregation. All the alleles were present in the haploid progeny. The presence of diverse alleles in embryos derived from the same anther demonstrated that they were induced on multiple microspores or pollen grains and they were not clonally propagated. Also, diploid cultures and mixtures of haploid-diploid tissues were obtained. The origin of such cultures, either somatic or gametic, was elucidated by SSR markers. All the embryos showed only one allele, corroborating a haploid origin. Allelic composition of the haploid progeny permitted parental identification among all analyzed trees.

Alleles↗
PubMed2003

Determining the role of N remobilization for growth of apple (Malus domestica Borkh) trees by measuring xylem-sap N flux.

The contribution of N remobilization to the seasonal growth of field-grown Malus domestica (apple) trees was measured using two different techniques. 'Fuji' trees grafted on M.9 apple rootstocks were planted in the field and fertilized and irrigated for two growing seasons. During the second year, the trees received 15N-labelled fertilizer and destructive harvests were taken during the spring and summer to determine the pattern of N remobilization and uptake. At the same time, patterns of N translocation in the xylem were measured by sampling saps at each harvest and analysing them for their constituent amino acids and amides. Total water flux through the trunk xylem was also measured throughout the sampling period using the heat balance technique. The flux of amino compounds in the xylem was then calculated to see if this approach could quantify remobilization. Most of the N for leaf growth was provided by remobilization, which lasted for some 40 d following bud-burst. The labelled N was not taken up until 14 d after remobilization had started. The predominant amino compounds recovered in the xylem were Asn, Asp, Arg, and Gln, whose concentration peaked during remobilization, except for Arg whose concentration was highest at bud-break and declined thereafter. The amount of N translocated in the xylem as Asn, Asp and Gln correlated well with the amount of N remobilized (as measured by the recovery of unlabelled N in the new above-ground growth). The data suggest that Arg is translocated predominantly as a consequence of root uptake and they are discussed in relation to measuring N remobilization in field-grown trees.

Amino Acids↗
PubMed1995

Phylogenetic test of the molecular clock and linearized trees.

To estimate approximate divergence times of species or species groups with molecular data, we have developed a method of constructing a linearized tree under the assumption of a molecular clock. We present two tests of the molecular clock for a given topology: two-cluster test and branch-length test. The two-cluster test examines the hypothesis of the molecular clock for the two lineages created by an interior node of the tree, whereas the branch-length test examines the deviation of the branch length between the tree root and a tip from the average length. Sequences evolving excessively fast or slow at a high significance level may be eliminated. A linearized tree will then be constructed for a given topology for the remaining sequences under the assumption of rate constancy. We have used these methods to analyze hominoid mitochondrial DNA and drosophilid Adh gene sequences.

Alcohol Dehydrogenase↗
PubMed2001

Tree stem diameter variations and transpiration in Scots pine: an analysis using a dynamic sap flow model.

A dynamic model for simulating water flow in a Scots pine (Pinus sylvestris L.) tree was developed. The model is based on the cohesion theory and the assumption that fluctuating water tension driven by transpiration, together with the elasticity of wood tissue, causes variations in the diameter of a tree stem and branches. The change in xylem diameter can be linked to water tension in accordance with Hookeâ s law. The model was tested against field measurements of the diurnal xylem diameter change at different heights in a 37-year-old Scots pine at Hyytiälä, southern Finland (61 degrees 51' N, 24 degrees 17' E, 181 m a.s.l.). Shoot transpiration and soil water potential were input data for the model. The biomechanical and hydraulic properties of wood and fine root hydraulic conductance were estimated from simulated and measured stem diameter changes during the course of 1 day. The estimated parameters attained values similar to literature values. The ratios of estimated parameters to literature values ranged from 0.5 to 0.9. The model predictions (stem diameters at several heights) were in close agreement with the measurements for a period of 6 days. The time lag between changes in transpiration rate and in sap flow rate at the base of the tree was about half an hour. The analysis showed that 40% of the resistance between the soil and the top of the tree was located in the rhizosphere. Modeling the water tension gradient and consequent woody diameter changes offer a convenient means of studying the link between wood hydraulic conductivity and control of transpiration.

Models, Biological↗
PubMed2001

Induction of flowering in tropical trees by a 30-min reduction in photoperiod: evidence from field observations and herbarium specimens.

During the late rainy season in October 1997 we observed. over a range of >100 km, the highly synchronous emergence of flower buds in several deciduous tree species of the semi-deciduous tropical forest in Guanacaste, Costa Rica. Synchronous flowering soon after the rapid decline in day length around the September equinox and in the absence of any notable climatic cues suggested flower induction by declining photoperiod. By combining field observations and the analysis of flowering herbarium collections, we established highly synchronous flowering periods with low interannual and latitudinal variation predicted for photoperiodic flower induction for more than 25 tree species and a few herbs. We describe morphogenetic changes at the shoot apex of three species during flower induction and the suppression and induction of flowering in several herbaceous species by experimental daylight extension. The combined observations provide strong, mainly indirect evidence for photoperiodic induction of flowering in many tropical tree species. At low latitudes with annual variation in day length of 1 hour, flower induction must be caused by a decline in photoperiod of 30 min or less. This is the first report of photoperiodic control of flowering in trees.

Costa Rica↗
PubMed2001

Photoperiodic control of seasonal development and dormancy in tropical stem-succulent trees.

Tropical stem-succulent trees store large quantities of water in their trunks yet remain leafless during the early and mid dry season. In contrast to most other tropical trees, bud break of vegetative buds is not induced in fully hydrated stem succulents between the winter solstice and the spring equinox by leaf abscission, abnormal rain showers or irrigation. Vegetative buds of leafless trees are therefore in a state of endo-dormancy similar to that of temperate perennial plants during early winter. Highly synchronous bud break regularly occurs soon after the spring equinox, often weeks before the first rainfalls of the wet season. These observations suggested that endo-dormancy and bud break might be induced by declining and increasing photoperiods after the autumn and spring equinoxes, respectively. In phenological field observations, we confirmed highly synchronous bud break after the spring equinox in many trees of five stem-succulent species in the northern and southern hemispheres. Shoot growth of potted saplings of Plumeria rubra L. was arrested by a decline in day length below 12 h after the autumn equinox, but continued in saplings maintained in a 13-h photoperiod. Conversely, exposure to a 13-h photoperiod induced bud break of dormant apical buds in saplings and cuttings in January, whereas plants maintained in the natural day length of < 11.7 h remained dormant. Photoperiodic control of endo-dormancy of vegetative buds in stem succulents is thus supported by field observations and experimental variation of the photoperiod. At low latitudes, where annual variation of day length is less than 1 h, bud dormancy is induced and broken by variations in photoperiod of less than 30 min.

Photoperiod↗
PubMed2002

Measurement carbon dioxide concentration does not affect root respiration of nine tree species in the field.

Inhibition of respiration has been reported as a short-term response of tree roots to elevated measurement CO2 concentration ([CO2]), calling into question the validity of root respiration rates determined at CO2 concentrations that differ from the soil [CO2] in the rooting zone. Our objectives were to validate previous observations of a direct CO2 effect on root respiration in sugar maple (Acer saccharum Marsh.) and to determine if high [CO2] also inhibited root respiration in other tree species. Root respiration rates for nine common North American tree species were measured in the field at ambient soil temperature at both 350 and 1000 microl CO2 l-1. No evidence of direct inhibition of root respiration by elevated measurement [CO2] was found for any of the species tested. The ratio of respiration rates at 1000 and 350 microl CO2 l-1 ranged from 0.97 to 1.07, and the 95% confidence intervals for this ratio included unity for all species tested. Tests of a respiration cuvette used in earlier experiments suggested that gas leakage from the cuvette/IRGA system created an apparent direct CO2 effect on respiration of sugar maple roots when none actually existed. Small sample masses used in those experiments exacerbated the error. Careful attention to the possibility of gas leaks and the avoidance of small sample masses should produce data that will allow researchers to accurately assess whether direct effects of measurement [CO2] exist. Our findings of no direct CO2 effect on respiration of roots of a wide variety of species suggest that such effects may be less common than previously thought for tree roots.

Carbon Dioxide↗
PubMed2002

Measured and predicted changes in tree and stand water use following high-intensity thinning of an 8-year-old Eucalyptus nitens plantation.

We investigated changes in the pattern of water use of an 8-year-old Eucalyptus nitens (Deane and Maiden) Maiden plantation soon after thinning. Sap flow sensors using heat pulse technology were deployed across three stands thinned to a final density of 100, 250 or 600 trees ha-1 plus an unthinned control (1250 trees ha-1). Changes in the relationship between tree size and daily water use were measured for 4 to 7 months after thinning. Thinning had no effect on sapwood water content. The increase in tree water use as a result of thinning was driven largely by significant changes in the radial pattern of sap velocity through the sapwood. The use of a canopy fraction factor in the Penman-Monteith equation to account for discontinuous canopies showed promise as a simple and effective method of scaling the model to predict transpiration from thinned plantations.

Circadian Rhythm↗
PubMed2002

Responses of deciduous broadleaf trees to defoliation in a CO2 enriched atmosphere.

Relatively little is known about the implications of atmospheric CO2 enrichment for tree responses to biotic disturbances such as folivory. We examined the combined effects of elevated CO2 concentration ([CO2]) and defoliation on growth and physiology of sugar maple (Acer saccharum Marsh.) and trembling aspen (Populus tremuloides Michx.). Seedlings were planted in the ground in eight open-top chambers. Four chambers were ventilated with CO2-enriched air (ambient + 283 micromol mol-1) and four chambers were supplied with ambient air. After 6 weeks of growth, half of the leaf area was removed on a subset of seedlings of each species in each CO2 treatment. We monitored subsequent biomass gain and allocation, along with leaf gas exchange and chemistry. Defoliation did not significantly affect final seedling biomass in either species or CO2 treatment. Growth recovery following defoliation was associated with increased allocation to leaf mass in maple and a slight enhancement of mean photosynthesis in aspen. Elevated [CO2] did not significantly affect aspen growth, and the observed stimulation of maple growth was significant only in mid-season. Correspondingly, simulated responses of whole-tree photosynthesis to elevated [CO2] were constrained by a decrease in photosynthetic capacity in maple, and were partially offset by reductions in specific leaf area and biomass allocation to foliage in aspen. There was a significant interaction between [CO2] and defoliation on only a few of the measured traits. Thus, the data do not support the hypothesis that atmospheric CO2 enrichment will substantially alter tree responses to folivory. However, our findings do provide further indication that regeneration-stage growth rates of certain temperate tree species may respond only moderately to a near doubling of atmospheric [CO2].

Acer↗
PubMed2002

Comparing the influence of site quality, stand age, fire and climate on aboveground tree production in Siberian Scots pine forests.

Temporal patterns of stem and needle production and total aboveground net primary production (ANPP) were studied at the tree and stand level along four chronosequences of Siberian Scots pine (Pinus sylvestris L.) forests differing in site quality (poor lichen type or the more fertile Vaccinium type) and in frequency of surface fires (unburned, moderately burned (fire return interval of approximately 40 years), or heavily burned (fire return interval of approximately 25 years)). The maximum range of variability in aboveground production was quantified for: (1) possible long-term changes in site quality; (2) stand age; (3) non-stand-replacing, recurring surface fires; and (4) interannual climate variability. For (1) and (2), total ANPP was low in the lichen-type chronosequence, reached a maximum of 170 g C m(-2) year(-1) after 100 years and decreased to 100 g C m(-2) year(-1) in older stands. Maximum ANPP in the Vaccinium-type chronosequence was 340 g C m(-2) year(-1) and occurred earlier in the 53-year-old stand than in the other stands. Along the lichen-type chronosequences, peak ANPP was paralleled by maximum carbon allocation to stem growth. (3) In mature trees, damage by recurrent surface fires decreased stem growth by 17 +/- 19% over a 10-year period relative to pre-fire values. At longer timescales, ANPP was hardly affected by fire-related differences in mortality. (4) Needle- plus stem-NPP, reconstructed for a 3-year period, varied within a range of 15 g C m(-2) year(-1) in the lichen-type stands and 35 g C m(-2) year(-1) in the Vaccinium-type stands. For the same period, the coefficient of variance was higher for needle-NPP (20 +/- 10%) than for stem-NPP (12 +/- 7%). Needle- and stem-NPP did not covary in time. Most 30-year time series of stem-NPP at the tree level exhibited strong autocorrelation. In older trees, stem-NPP was positively correlated with growing season precipitation. Thus, the factors driving variability in ANPP ranked according to their maximum influence as: stand age (controlled by the frequency of stand-replacing fires) > site quality > growth depression because of surface fire damage approximately equal age-related reduction in ANPP > interannual variability approximately equal long-term effects of fire (stand density reduction). In lichen-type forests, we found that ANPP at the landscape level declined sharply when the interval between stand-replacing fires was less than 120 years, illustrating that fire strongly influences ANPP of boreal Scots pine forests.

Climate↗
PubMed2003

Internal oxygen transport in cuttings from flood-adapted várzea tree species.

Tree species from the Central Amazon inundation areas are subjected to extreme flooding, with trees being partially submerged for up to 10 months. The rapidly advancing floodwater table at the onset of the aquatic phase interrupts the inward diffusion of oxygen from the atmosphere to submerged plant parts. Salix martiana (Leyb.) (Salicaceae) and Tabernaemontana juruana ((Markgr.) Schumann ex J. F. Macbride) (Apocynaceae), tree species typical of Amazon floodplains, respond to low oxygen concentrations by forming adventitious roots capable of longitudinal oxygen transport. Cuttings of these tree species were subjected to simulated flooding and the oxygen concentration of the root cortex was temporally monitored by oxygen microelectrodes that penetrated the roots. Changes in the floodwater table made it possible to localize precisely the entry points of atmospheric oxygen. Under experimental conditions, mathematical description of the transport kinetics revealed that longitudinal transport of O2 in both species was mainly attributable to diffusion. Based on the finding that diffusion was inhibited by a small increase in the floodwater table, we conclude that internal oxygen transport during a rising water table is only attainable when adventitious roots are continuously and rapidly developed, as is the case in S. martiana. In T. juruana, slow growth of adventitious roots and low root porosity suggest that other adaptations are required to overcome long flooding periods.

Adaptation, Physiological↗
PubMed2004

Winter variation in xylem sap pH of walnut trees: involvement of plasma membrane H+-ATPase of vessel-associated cells.

We studied seasonal variation in xylem sap pH of Juglans regia L. Our main objectives were to (1) test the effect of temperature on seasonal changes in xylem sap pH and (2) study the involvement of plasma membrane H+-ATPase of vessel-associated cells in the control of sap pH. For this purpose, orchard-grown trees were compared with trees grown in a heated (> or = 15 degrees C) greenhouse. During autumn, sap pH was not directly influenced by temperature. A seasonal change in H+-ATPase activity resulting from seasonal variation in the amount of protein was measured in orchard-grown trees, whereas no significant seasonal changes were recorded in greenhouse-grown trees. Our data suggest that H+-ATPase does not regulate xylem sap pH directly by donating protons to the xylem, but by facilitating secondary active H+/sugar transport, among other mechanisms.

Hydrogen-Ion Concentration↗
PubMed2004

Interstock-induced mechanism of increased growth and salt resistance of orange (Citrus sinensis) trees.

Interstocks improve the growth and salt resistance of lemon (Citrus limon (L.) Burm. f.) trees, but their effects on orange (Citrus sinensis (L.) Osbeck) trees are unknown. We grew 'Cleopatra' mandarin (CM) seedlings, budded trees of 'Salustiano' orange (SAO) on CM, 'Valencia Late' orange (VLO) on CM (VLO/CM), and interstock trees VLO/SAO/CM in pots of sand watered with nutrient solution containing 5 (control) or 50 mM NaCl for 12 weeks. Plants were harvested on six successive occasions and the time trends in relative growth rate (RGR) and its components were estimated by fitting a Richards function regression to the harvest data. At low and high salinities, the VLO/SAO/CM combination had higher mean RGR than VLO/CM. Under control conditions, the increase in RGR caused by the interstock was the result of an increase in leaf mass fraction (LMF; leaf dry mass/plant dry mass ratio). Increases in net assimilation rate on a leaf mass basis (NARm) and LMF contributed equally to the increase in RGR in saline conditions, their growth response coefficients being 0.52 and 0.48, respectively. The structural modifications, specific leaf area (SLA) and leaf area ratio (LAR; leaf area:plant dry mass ratio), had a slight influence on the reduction in RGR by salinity. However, NARm had a large influence on RGR, except in CM. The interstock-induced mechanism increased biomass allocation to the assimilatory organs and, under saline conditions, increased Cl- and Na+ allocations to roots. Thus, the flux of ions to the leaves was either delayed or reduced or both. The dilution of imported ions by foliar growth reduced ion concentrations in leaves, resulting in higher NARm, which together with higher LMF, increased RGR.

Citrus sinensis↗
PubMed2005

Modeling tree water deficit from microclimate: an approach to quantifying drought stress.

Tree water deficit estimated by measuring water-related changes in stem radius (DeltaW) was compared with tree water deficit estimated from the output of a simple, physiologically reasonable model (DeltaWE), with soil water potential (Psisoil) and atmospheric vapor pressure deficit (VPD) as inputs. Values of DeltaW were determined by monitoring stem radius changes with dendrometers and detrending the results for growth. We followed changes in DeltaW and DeltaWE in Pinus sylvestris L. and Quercus pubescens Willd. over 2 years at a dry site (2001-2002; Salgesch, Wallis) and in Picea abies (L.) Karst. for 1 year at a wet site (1998; Davos, Graubuenden) in the Swiss Alps. The seasonal courses of DeltaW in deciduous species and in conifers at the same site were similar and could be largely explained by variation in DeltaWE. This finding strongly suggests that DeltaW, despite the known species-specific differences in stomatal response to microclimate, is mainly explained by a combination of atmospheric and soil conditions. Consequently, we concluded that trees are unable to maintain any particular DeltaW. Either Psisoil or VPD alone provided poorer estimates of DeltaW than a model incorporating both factors. As a first approximation of DeltaWE, Psisoil can be weighted so that the negative mean Psisoil reaches 65 to 75% of the positive mean daytime VPD over a season (Q. pubescens: approximately 65%, P. abies: approximately 70%, P. sylvestris: approximately 75%). The differences in DeltaW among species can be partially explained by a different weighting of Psisoil against VPD. The DeltaW of P. sylvestris was more dependent on Psisoil than that of Q. pubescens, but less than that of P. abies, and was less dependent on VPD than that of P. abies and Q. pubescens. The model worked well for P. abies at the wet site and for Q. pubescens and P. sylvestris at the dry site, and may be useful for estimating water deficit in other tree species.

Climate↗
PubMed2005

Stomatal and non-stomatal limitations to photosynthesis in four tree species in a temperate rainforest dominated by Dacrydium cupressinum in New Zealand.

We assessed the relative limitations to photosynthesis imposed by stomatal and non-stomatal processes in Dacrydium cupressinum Lamb. (Podocarpaceae), which is the dominant species in a native, mixed conifer-broad-leaved rainforest in New Zealand. For comparison, we included three co-occurring broad-leaved tree species (Meterosideros umbellata Cav. (Myrtaceae), Weinmannia racemosa L.f. (Cunoniaceae) and Quintinia acutifolia Kirk (Escalloniaceae)) that differ in phylogeny and in leaf morphology from D. cupressinum. We found that low foliage phosphorus content on an area basis (P(a)) limited light-saturated photosynthesis on an area basis (A(sat)) in Q. acutifolia. Depth in the canopy did not generally affect A(sat) or the relative limitations to A(sat) because of stomatal and non-stomatal constraints, despite reductions in the ratio of foliage mass to area, foliar nitrogen on an area basis (N(a)) and P(a) with depth in the canopy. In the canopy-dominant conifer D. cupressinum, A(sat) was low, consistent with low values of the maximum rate of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylation (V(cmax)). In comparison, the A(sat) response of the three broad-leaved tree species was quite variable. Although A(sat) was high in the canopy-dominant M. umbellata, it was low in the sub-canopy trees W. racemosa and Q. acutifolia. Relative stomatal limitation to photosynthesis was more pronounced in W. racemosa (40%) than in the other three species (28-33%). Despite differences in degree, non-stomatal limitation to A(sat) predominated in all tree species.

Chlorophyll↗
PubMed2005

Bridging process-based and empirical approaches to modeling tree growth.

The gulf between process-based and empirical approaches to modeling tree growth may be bridged, in part, by the use of a common model. To this end, we have formulated a process-based model of tree growth that can be fitted and applied in an empirical mode. The growth model is grounded in pipe model theory and an optimal control model of crown development. Together, the pipe model and the optimal control model provide a framework for expressing the components of tree biomass in terms of three standard inventory variables: tree height, crown height and stem cross-sectional area. Growth rates of the inventory variables and the components of biomass are formulated from a carbon balance. Fundamentally, the parameters of the model comprise physiological rates and morphological ratios. In principle, the values of these parameters may be estimated by lower-level process models. Alternatively, the physiological and morphological parameters combine, under reasonable assumptions, into a set of aggregate parameters, whose values can be estimated from inventory data with a statistical fitting procedure.

Biomass↗
PubMed2005

Modeling the dynamics of pressure propagation and diameter variation in tree sapwood.

A non-steady-state model of water tension propagation in tree stems was developed. The model is based on the cohesion theory and the assumption that fluctuating water tension driven by transpiration together with the elasticity of wood cause variations in the diameter of a tree stem. The change in xylem diameter can be linked to water tension in accordance with Hooke's law. The model was tested against field measurements of the diurnal change in xylem diameter at different heights in a 180-year-old Scots pine tree at Hyytiälä, southern Finland. Model predictions agreed well with measurements. The effect of tree dimensions on pressure propagation was examined with the model. The model outcomes were also consistent with results of several field measurements presented in the literature.

Models, Biological↗
PubMed2006

Mechanical and ecophysiological significance of the form of a young Acer rufinerve tree: vertical gradient in branch mechanical properties.

Most tree biomechanics models assume uniformity of mechanical properties within a tree, and only a few studies have focused on differences in mechanical status among branches. We examined mechanical properties of 49 branches of two 10-year-old trees of Acer rufinerve Sieb. et Zucc. For each branch, bending moment due to its own fresh mass, elastic modulus, section modulus and flexural stiffness were obtained. Elastic modulus of the branch was correlated with the density and thickness of the fiber cell wall and decreased with crown depth, indicating that branches at lower positions were more elastic than branches at upper positions. Compared to lower branches, upper branches were less inclined, possessed thicker growth rings, more long shoots and were subject to smaller stresses. The leaf arrangement in the upper branches might be effective in transmitting more light to the lower branches. In contrast, the lower branches were more inclined toward the horizontal and subject to greater stresses than the upper branches. Lower branch inclinations were attributed to smaller dry matter investment in diameter growth. Upper and lower branch inclinations were slightly greater and smaller, respectively, than those predicted by beam theory. The alleviation in inclination of the lower branches is probably caused by negative gravitropic responses such as tension wood formation or upward shoot elongation, or both. The horizontal display of leaves in the lower branches would be effective in light interception. The reduction in cost of the lower branches can be adaptive because they have a shorter life expectancy than the upper branches. The results showed that an adaptive tree form is realized by a vertical gradient in branch mechanical properties.

Acer↗