Genetic variation in foliar nutrient concentration in relation to foliar carbon isotope composition and tree growth with clones of the F1 hybrid between slash pine and Caribbean pine

The objectives of this study were: (1) to quantify the genetic variation in foliar nutrient concentration in relation to foliar carbon isotope composition (δ¹³C) and tree growth of 122 clones of ca. 4-year-old F₁ hybrids between slash pine (Pinus elliottii Engelm var. elliottii) and Caribbean pine (P. caribaea var. hondurensis Barr. et Golf.) grown at two experimental sites with different water and nutrient availability in southeast Queensland, Australia and (2) to examine the potential of using foliar nutrient concentration of the 4-year-old tree canopies for selecting elite F₁ hybrid pine clones with improved nutrient-use efficiency (NUE) and water-use efficiency (WUE), and ultimately enhanced tree growth under ambient growing conditions. There were significant differences in foliar nutrient concentrations between two canopy positions (upper outer and lower outer canopy) sampled, between summer and winter, and between the two sites. This highlights that foliar nutrient concentrations are influenced by sampling and environment. Significant genetic variations in foliar nutrient concentrations were detected between the clones, between the female parents, and between the male parents of the clones in both sampling seasons at both sites. Depending on the nutrient concerned, canopy position, season, and site sampled, the clones accounted for 4.7–33.9% of the total variation in foliar nutrient concentrations, the clone female parents for 0–25.1% and the clone male parents for 0–28.6%. The site-by-clone interactions were statistically significant for foliar N, P, Mg, Cu, Zn, Mn, Fe and mineral concentrations at the upper outer canopy in summer, and for foliar N concentration in winter. There were significant, positive correlations between clone means of foliar δ¹³C and N concentration at the upper outer canopy in summer for the wet site, while clone foliar δ¹³C was also positively related to clone foliar N concentration at both canopy positions in summer for the dry site. This suggests that clone WUE as reflected in foliar δ¹³C may be improved by selecting elite clones with higher foliar N concentration and increased photosynthesis, leading to enhanced tree growth when both water and N are the major growth-limiting factors. This is supported by the positive correlation detected between clone tree height and foliar N concentration at the upper outer canopy for both sites. Thus, foliar nutrient (particularly N) concentration, together with foliar δ¹³C, may be useful for assisting in selection of exotic pine clones with improved NUE and WUE, and enhanced tree growth under the nutrient- and water-limiting environments.     

Branchlet nutrient concentration in hoop pine (Araucaria cunninghamii) relative to family,stable carbon and oxygen isotope ratios and growth rate in contrasting environments

Genetic variation in branchlet nutrient (N, P, K, Na, Ca, Mg, Mn and Fe) concentrations and mineral concentration (sum of branchlet P, K, Na, Ca, Mg, Mn and Fe concentrations) of 8-9-year-old hoop pine (Araucaria cunninghamii Ait. ex D. Don) half-sib families was assessed for four canopy positions at a wet site (23 families) and two canopy positions at an N- and water-limiting dry site (22 families) in relation to tree growth and associated branchlet carbon (delta13C) and oxygen (delta18O) isotope composition in southeast Queensland, Australia. Branchlet nutrient and mineral concentrations varied significantly among families and with canopy position and site. Depending on the canopy position sampled, the hoop pine family effect accounted for 0 to 13.8% of the total variation in branchlet N concentration, and for 0 to 30.3% of the total variation in branchlet mineral concentration at the wet site. The corresponding values for the family effect at the dry site were 0-13.3% for branchlet N concentration and 0-25.7% for branchlet mineral concentration. There were significant variations in branchlet P, K, Ca and Mg concentrations at both sites, and these variations differed with canopy position. Relationships between family means of branchlet N concentration and tree growth or delta13C or delta18O varied with canopy position at both sites. At the wet site, there were significant positive correlations between branchlet mineral concentration in the upper-outer or upper-inner canopy and tree height (r = 0.26 and 0.37, P < 0.01) and between branchlet mineral concentration and delta13C (r = 0.24, P < 0.01) in the upper-inner canopy, and a significant negative correlation between branchlet mineral concentration and delta13C (r = -0.21, P < 0.05) in the upper-outer canopy. At the dry site, branchlet mineral concentrations in the upper-inner and upper-outer canopy were significantly correlated with branchlet delta13C (r = -0.28 and -0.51, P < 0.01), and branchlet N concentration in the upper-inner canopy was significantly correlated with tree growth (r = 0.29, P < 0.01). A significant correlation between branchlet delta18O (an index of stomatal conductance) and branchlet mineral concentration at the dry site (r = 0.39, P = 0.020) indicated that stomatal conductance might be a factor regulating the variation in branchlet mineral concentration of the hoop pine families. Both branchlet N concentration and mineral concentration at particular canopy positions assist in selecting hoop pine families with improved tree growth and N- and water-use efficiency in environments where both N deficiency and a limited water supply are major factors affecting plantation productivity.     

Foliar delta(13)C and delta(18)O reveal differential physiological responses of canopy foliage to pre-planting weed control in a young spotted gum (Corymbia citriodora subsp. Variegata) plantation

Weed control may improve the growth of forest plantations by influencing soil water and nutrient availability, but our knowledge of leaf-level physiological responses to weed control at different within-canopy positions is limited for tropical and subtropical plantations. Foliar carbon (delta(13)C) and oxygen (delta(18)O) isotope compositions, gas exchange, and nitrogen (N(mass)) and phosphorus (P(mass)) concentrations at four canopy positions were assessed in a young spotted gum (Corymbia citriodora subsp. Variegata (F. Muell.) A.R. Bean & M.W. McDonald) plantation subjected to either weed control or no weed control treatment, to test if leaves at different positions within the tree canopy had the same physiological responses to the weed control treatment. Weed control increased foliar delta(13)C but lowered delta(18)O in the upper-outer and upper-inner canopy, indicating that weed control resulted in a higher foliar photosynthetic capacity at upper-canopy positions, a conclusion confirmed by gas exchange measurements. The increased photosynthetic capacity resulting from weed control can be explained by an increase in foliar N(mass). In the lower-outer canopy, weed control reduced foliar delta(13)C while lowering delta(18)O even more than in the upper-canopy, suggesting strong enhancement of the partial pressure of CO(2) in the leaf intercellular spaces and of foliar stomatal conductance in lower-canopy foliage. This conclusion was supported by gas exchange measurements. Foliar photosynthesis in the lower-inner canopy showed no significant response to weed control. The finding that leaves at different canopy positions differ in their physiological responses to weed control highlights the need to consider the canopy position effect when examining competition for soil nutrient and water resources between weeds and trees.     

Variation in branchlet delta13C in relation to branchlet nitrogen concentration and growth in 8-year-old hoop pine families (Araucaria cunninghamii) in subtropical Australia

Carbon isotope composition (delta13C) of branchlet tissue at nine canopy positions, and nitrogen concentration (N(mass)) at four canopy positions, were assessed in 8-year-old hoop pine (Araucaria cunninghamii Ait. ex D. Don) trees from 23 half-sib families, grown in six blocks of a progeny test in southeastern Queensland, Australia. There was considerable variation among sampling positions, families and blocks in both delta13C and N(mass). The delta13C was positively related to N(mass) only for samples from the upper outer crown (P < 0.005). Phenotypic correlations existed between tree growth and canopy delta13C. Branchlet delta13C of the inner and lower outer crown was positively related (P < 0.037) to tree height, but delta13C in branchlets of the upper outer crown was not related to tree height, or was related negatively (P < 0.045). There were significant differences in delta13C between hoop pine families for six canopy positions (upper canopy positions as well as lower canopy positions on the northern side), with heritabilities greater than 0.40. The significance of these findings is discussed in relation to water and light competition within the tree canopy of hoop pine.     

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 A?t.) 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.     

Co-occurring species differ in tree-ring delta(18)O trends

The stable oxygen isotope ratio (delta(18)O) of tree-ring cellulose is jointly determined by the delta(18)O of xylem water, the delta(18)O of atmospheric water vapor, the humidity of the atmosphere and perhaps by species-specific differences in leaf structure and function. Atmospheric humidity and the delta(18)O of water vapor vary seasonally and annually, but if the canopy atmosphere is well mixed, atmospheric characteristics should be uniform among co-occurring trees. In contrast, xylem water delta(18)O is determined by the delta(18)O of water being drawn from the soil, which varies with depth. If co-occurring trees draw water from different soil depths, this soil-water delta(18)O signal would be manifest as differences in delta(18)O among the trees. We examined the variation in tree ring delta(18)O, over eight decades during the 20th Century, among three species co-occurring in natural forest stands of the northern Rocky Mountains in the USA. We sampled 10 Douglas-firs (Pseudotsuga menziesii (Mirb.) Franco var. glauca), 10 ponderosa pines (Pinus ponderosa Laws.) and seven western white pines (Pinus monticola Dougl.). As expected, variation in atmospheric conditions was recorded in the delta(18)O of the cellulose produced in a given year, but observed climatic correlations with delta(18)O were weak. Significant correlations with June climate data included: daily maximum temperature (r = 0.29), daily minimum temperature (r = -0.25), mean temperature (r = 0.20), mean daily precipitation (r = -0.54), vapor pressure deficit (r = 0.32) and solar radiation (r = 0.44). Lagged effects were observed in Douglas-fir and western white pine. In these species, the delta(18)O of a given annual ring was correlated with the delta(18)O of the previous ring. Ponderosa pine showed no significant autocorrelation. Although the species means were correlated among years (r = 0.67 to 0.76), ponderosa pine was consistently enriched in delta(18)O relative to the other species; differences were close to 2 per thousand and they are steadily increasing. Relative to the mean for the three species, ponderosa pine is becoming steadily more enriched (-1.0 per thousand). In contrast, Douglas-fir is being steadily depleted and western pine is intermediate, with an enrichment of 0.5 per thousand. Because all trees were exposed to the same atmospheric conditions, the differences in delta(18)O observed between species are likely due either to differences in the depth of water extraction or leaf function. If the former, presumably ponderosa pine has steadily taken up more water from near the soil surface and Douglas-fir has shifted uptake to a greater depth. If the latter, we suggest the pronounced changes in leaf-water delta(18)O are a result of changes in leaf structure and function with tree size and age.     

Analyses of delta(13)C and delta(18)O in tree rings of Callitris columellaris provide evidence of a change in stomatal control of photosynthesis in response to regional changes in climate

We examined relationships between stable isotopes of carbon (delta(13)C) and oxygen (delta(18)O) in tree rings of Callitris columellaris F. Muell in the semi-arid Pilbara region of north-western Australia. To test the hypothesis that stomatal control of photosynthesis decreases during drier periods, we developed delta(13)C and delta(18)O chronologies spanning 1919-1999, and used a permutation regression approach to relate a 21-year running correlation between delta(13)C and delta(18)O to rainfall and temperature at Marble Bar and our study site. The relationship between delta(13)C and delta(18)O switched from being always negative before 1955 to being consistently positive after 1976, suggesting an increase in stomatal control of photosynthesis in recent decades. Changes in the delta(13)C-delta(18)O relationship reflected changes in rainfall, which has increased in the region by 30% since 1976. The correlation between delta(13)C and delta(18)O was positively related to the 21-year running mean of normalized rainfall anomalies at both the study site (P = 0.045, Adj. r(2) = 0.47) and Marble Bar (P = 0.046, Adj. r(2) = 0.48). In addition, the delta(13)C-delta(18)O correlation was negatively related (P = 0.047, Adj. r(2) = 0.61) to temperatures at Marble Bar. Our interpretation of the role of changes in climate affecting the relationship between tree-ring delta(13)C and delta(18)O is supported by evidence from the isotope composition of foliage samples: foliar delta(13)C and delta(18)O were negatively correlated with log stomatal conductance (delta(13)C, r = -0.41; delta(18)O, r = -0.42), whereas the correlation between foliar delta(13)C and delta(18)O was positive (r = 0.63, P = 0.027) after the summer wet period. Our data indicate that stomatal control of photosynthesis in Callitris adjusts to region-wide changes in climate and that, in a warmer and drier world, trees might adapt by increasing non-stomatal control of photosynthesis.     

Boxelder water sources and physiology at perennial and ephemeral stream sites in Arizona

To assess the influence of stream water on leaf gas exchange and water potential in different sized boxelder trees (Acer negundo L.), we compared these characteristics in trees growing beside a perennial stream and a nearby ephemeral stream in a montane-riparian forest in northern Arizona. Patterns of tree water use were quantified by stable isotope analysis (delta(18)O). Physiological characteristics were similar for large and small trees. Similarity between sites in predawn and daytime water potentials and xylem delta(18)O indicated that stream water was not a physiologically important water source. Seasonal and site variations in light-saturated net photosynthetic rate were significantly related to leaf-to-air vapor pressure deficit (r = -0.691) and foliar nitrogen concentration (r = 0.388). Although deep water was the dominant water source, surface soil water was utilized following precipitation, especially by small trees. We conclude that net carbon gain and severity of water stress are only weakly coupled to stream water availability.     

Loblolly pine growth and soil nutrient stocks eight years after forest slash incorporation

Incorporation of forest slash during stand establishment is proposed as a means of increasing soil carbon and nutrient stocks. If effective, the increased soil carbon and nutrient status may result in increased aboveground tree growth. Eight years after study installation, the impact of forest slash incorporation into the soil on soil carbon and nutrient stocks, foliar nutrients and loblolly pine growth are examined on mineral and organic sites on the North Carolina Lower Coastal Plain. Treatments include leaving forest slash on the surface and flat planting (control); V-shear and bedding (conventional), mulch forest slash followed by bedding (strip mulch) and mulch forest slash and till into the soil followed by bedding (strip mulch till). After eight years, mulching and/or tillage did not have a significant impact (p > 0.05) on soil bulk density or soil chemical properties (pH, cation exchange capacity, soil nutrients). Additionally, neither tree foliar nutrients nor stand volume were significantly impacted. However, significant effects were observed for soil phosphorus contents and stand volume between the control plots and the other treatment plots. For example, the mean stand volumes on the mineral site were 24.49 ± 1.28, 38.16 ± 2.90, 44.59 ± 3.07 and 46.96 ± 2.74 m³ ha⁻¹ for the control, conventional, strip mulch and strip mulch till plots. These observations are more likely due to the effect of bedding rather than mulching or tillage of the forest slash. These results are consistent for the mineral and the organic sites. Considering the greater expense to install the mulch and tillage treatments, the lack of a treatment effect on soil carbon and nutrient stocks and tree growth does not justify these treatments on these sites.     

Genetic effects on total phenolics, condensed tannins and non-structural carbohydrates in loblolly pine (Pinus taeda L.) needles

Carbon allocation to soluble phenolics (total phenolics, proanthocyanidins (PA)) and total non-structural carbohydrates (TNC; starch and soluble sugars) in needles of widely planted, highly productive loblolly pine (Pinus taeda L.) genotypes could impact stand resistance to herbivory, and biogeochemical cycling in the southeastern USA. However, genetic and growth-related effects on loblolly pine needle chemistry are not well characterized. Therefore, we investigated genetic and growth-related effects on foliar concentrations of total phenolics, PA and TNC in two different field studies. The first study contained nine different genotypes representing a range of genetic homogeneity, growing in a 2-year-old plantation on the coastal plain of North Carolina (NC), USA. The second study contained eight clones with different growth potentials planted in a 9-year-old clonal trial replicated at two sites (Georgia (GA) and South Carolina (SC), USA). In the first study (NC), we found no genetic effects on total phenolics, PA and TNC, and there was no relationship between genotype size and foliar biochemistry. In the second study, there were no differences in height growth between sites, but the SC site showed greater diameter (diameter at breast height (DBH)) and volume, most likely due to greater tree mortality (lower stocking) which reduced competition for resources and increased growth of remaining trees. We found a significant site?×?clone effect for total phenolics with lower productivity clones showing 27-30% higher total phenolic concentrations at the GA site where DBH and volume were lower. In contrast to the predictions of growth-defense theory, clone volume was positively associated with total phenolic concentrations at the higher volume SC site, and PA concentrations at the lower volume GA site. Overall, we found no evidence of a trade-off between genotype size and defense, and genetic potential for improved growth may include increased allocation to some secondary metabolites. These results imply that deployment of more productive loblolly pine genotypes will not reduce stand resistance to herbivory, but increased production of total phenolics and PA associated with higher genotype growth potential could reduce litter decomposition rates and therefore, nutrient availability.     

Diurnal changes in gas exchange and carbon partitioning in needles of fast- and slow-growing families of loblolly pine (Pinus taeda)

We investigated diurnal and seasonal changes in carbon acquisition and partitioning of recently assimilated carbon in fast- and slow-growing families of loblolly pine (Pinus taeda L.) to determine whether fast-growing families exhibited greater carbon gain at the leaf level. Since planting on a xeric infertile site in Scotland County, NC, USA in 1993, five Atlantic Coastal Plain (ACP) and five "Lost Pines" Texas (TX) families have been grown with either optimal nutrition or without fertilization (control). In 1998 and 1999, gas exchange parameters were monitored bimonthly in four families and needles were analyzed bimonthly for starch and soluble sugar concentrations. Although diurnal and seasonal effects on net photosynthesis (A(net)) and maximum rate of light-saturated photosynthesis (A(max)) were significant, few family or treatment differences in gas exchange characteristics were observed. The A(net) peaked at different times during the day over the season, and A(max) was generally highest in May. Instantaneous water-use efficiency (WUE(i)), derived from gas exchange parameters, did not differ among families, whereas foliage stable isotope composition (delta(13)C) values suggested that TX families exhibited lower WUE than more mesic ACP families. Although there were no diurnal effects on foliar starch concentrations, needles exhibited pronounced seasonal changes in absolute concentrations of total nonstructural carbohydrates (TNC), starch and soluble sugars, and in partitioning of TNC to starch and sugars, mirroring seasonal changes in photosynthesis and shoot and root growth. In all families, foliar starch concentrations peaked in May and decreased to a minimum in winter, whereas reducing sugar concentrations were highest in winter. Some family and treatment differences in partitioning of recently assimilated carbon in needles were observed, with the two TX families exhibiting higher concentrations of TNC and starch and enhanced starch partitioning compared with the ACP families. We conclude that growth differences among the four families are not a function of differences in carbon acquisition or partitioning at the leaf level.     

Variations in growth, survival and carbon isotope composition (delta(13)C) among Pinus pinaster populations of different geographic origins

To evaluate differences in growth and adaptability of maritime pine (Pinus pinaster Ait.), we studied growth, polycyclism, needle tissue carbon isotope composition (delta(13)C) as an estimate of water-use efficiency (WUE) and survival of seven populations at 10 years of age growing in a performance trial at a provenance test site in Escaroupim, Portugal. Six populations were from relatively high rainfall sites in Portugal and southwestern France (Atlantic group), and one population was from a more arid Mediterranean site in Spain. There were significant differences between some populations in total height, diameter at breast height, delta(13)C of bulk needle tissue, polycyclism and survival. A population from central Portugal (Leiria, on the Atlantic coast) was the tallest and had the lowest delta(13)C. Overall, the variation in delta(13)C was better explained by the mean minimum temperatures of the coldest month than by annual precipitation at the place of origin. Analyses of the relationships between delta(13)C and growth or survival revealed a distinct pattern for the Mediterranean population, with low delta(13)C (and WUE) associated with the lowest growth potential and reduced survival. There were significant negative correlations between delta(13)C and height or survival in the Atlantic group. Variation in polycyclism was correlated with annual precipitation at the place of origin. Some Atlantic populations maintained a high growth potential while experiencing moderate water stress. A detailed knowledge of the relationships between growth, survival and delta(13)C in contrasting environments will enhance our ability to select populations for forestry or conservation.     

Dry mass allocation, water use efficiency and delta(13)C in clones of Eucalyptus grandis, E. grandis x camaldulensis and E. grandis x nitens grown under two irrigation regimes

Clonal variation in water use efficiency (WUE), dry mass accumulation and allocation, and stable carbon isotope ratio (delta(13)C) of crude leaf fiber extracts was determined in six clones of Eucalyptus grandis W. Hill ex Maiden. grown for 16 months in field lysimeters in two soil water regimes. The relationships between delta(13)C and WUE calculated on the basis of leaf, harvestable stem, shoot and whole-plant dry mass accumulation were investigated. There was no clonal variation in dry mass accumulation but clonal allocation to roots, harvestable stems, branches and leaves varied. Water use efficiencies (mass of plant or plant part/water used over 16 months) differed significantly between clones. The clonal ranking of WUE varied depending on the units of dry mass accumulation used. Significant relationships between delta(13)C values and instantaneous water use efficiencies and ratios of internal leaf to ambient CO(2) concentrations were found only in the high soil water treatment. There were no relationships between delta(13)C values and whole-plant, shoot and harvestable stem water use efficiencies and soil water availability. Values of delta(13)C were negatively correlated with dry mass accumulation in the low soil water treatment. At the whole-plant level, WUE was positively correlated with dry mass accumulation in the high soil water treatment. We found significant differences in delta(13)C values between clones and the clonal rankings in delta(13)C and WUE were maintained in both soil water treatments.     

Regulation of ponderosa pine foliar physiology and insect resistance mechanisms by basal area treatments

We compared foliar physiology and several measures of tree resistance to insect attack among ponderosa pine (Pinus ponderosa var. scopulorum Engelm.) trees growing in thinned stands. Measurements were made in a second-growth ponderosa pine forest in northern Arizona where the basal area treatments (6.9, 18.4, 27.6, 78.2 m(2) ha(-1)) have been experimentally maintained by frequent thinnings for 32 years before our measurements began in 1994. Most of the physiological characteristics measured were affected by the basal area treatments. As stand basal area increased from 6.9 to 78.2 m(2) ha(-1), predawn water potential, midday water potential, net photosynthetic rate, resin production, phloem thickness, and foliar toughness decreased. Foliar nitrogen concentration was greatest in trees in the intermediate basal area treatments. Our results show that the physiological condition of second-growth ponderosa pine can be manipulated by silvicultural control of stand basal area, and support the hypothesis that high stand basal area increases tree stress and decreases tree resistance to insect attack.     

Element accumulation patterns of deciduous and evergreen tree seedlings on acid soils: implications for sensitivity to manganese toxicity

Foliar nutrient imbalances, including the hyperaccumulation of manganese (Mn), are correlated with symptoms of declining health in sensitive tree species growing on acidic forest soils. The objectives of this study were to: (1) compare foliar nutrient accumulation patterns of six deciduous (sugar maple (Acer saccharum Marsh.), red maple (Acer rubrum L.), red oak (Quercus rubra L.), white oak (Quercus alba L.), black cherry (Prunus serotina Ehrh.) and white ash (Fraxinus americana L.)) and three evergreen (eastern hemlock (Tsuga canadensis L.), white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss.)) tree species growing on acidic forest soils; and (2) examine how leaf phenology and other traits that distinguish evergreen and deciduous tree species influence foliar Mn accumulation rates and sensitivity to excess Mn. For the first objective, leaf samples of seedlings from five acidic, non-glaciated field sites on Pennsylvania's Allegheny Plateau were collected and analyzed for leaf element concentrations. In a second study, we examined growth and photosynthetic responses of seedlings exposed to excess Mn in sand culture. In field samples, Mn in deciduous foliage hyperaccumulated to concentrations more than twice as high as those found in evergreen needles. Among species, sugar maple was the most sensitive to excess Mn based on growth and photosynthetic measurements. Photosynthesis in red maple and red oak was also sensitive to excess Mn, whereas white oak, black cherry, white ash and the three evergreen species were tolerant of excess Mn. Among the nine species, relative rates of photosynthesis were negatively correlated with foliar Mn concentrations, suggesting that photosynthetic sensitivity to Mn is a function of its rate of accumulation in seedling foliage.     

The effects of ultraviolet-B radiation on photosynthesis of different aged needles in field-grown loblolly pine

We examined the effect of supplemental UV-B radiation (290-320 nm) on photosynthetic characteristics of different aged needles of 3-year-old, field-grown loblolly pine (Pinus taeda L.). Needles in four age classes were examined: I, most recently fully expanded, year 3; II, first flush, year 3; III, final flush, year 2; and IV, oldest needles still present, year 2. Enhanced UV-B radiation caused a statistically significant decrease (6%) in the ratio of variable to maximum fluorescence (F(v)/F(m)) following dark adaptation only in needles from the youngest age class, suggesting transient damage to photosynthesis. However, no effects of enhanced UV-B radiation on other instantaneous measures of photosynthesis, including maximum photosynthesis, apparent quantum yield and dark respiration, were seen for needles of any age. Foliar nitrogen concentration was unaffected by UV-B treatment. However, the (13)C/(12)C carbon isotope ratios (delta(13)C-a time integrated measure of photosynthetic function) of needles in age classes II and IV were 3% (P < 0.01) and 2% (P < 0.05) more negative, respectively, in treated plants than in control plants. Exposure to enhanced UV-B radiation caused a 20% decrease in total biomass and a 4% (P < 0.05), 25% (P < 0.01), and 9% (P < 0.01) decrease in needle length of needles in age classes I, II, and IV, respectively. The observed decreases in delta(13)C, and F(v)/F(m) of the needles in the youngest needle age class suggest subtle damage to photosynthesis, although overall growth reductions were probably a result of decreased total leaf surface rather than decreased photosynthetic capacity. Needles of age class IV had lower light- and CO(2)-saturated maximum photosynthetic rates (39%), lower dark respiration (34%), lower light saturation points (37%), lower foliar nitrogen concentration (28%), and lower delta(13)C (14%) values than needles of age class I. Apparent quantum yield and F(v)/F(m) did not change with needle age. The observed changes in photosynthesis and foliage chemical composition with needle age are consistent with previous studies of coniferous trees and may represent adaptations of older needles to shaded conditions within the canopy.     

Replacement of wildfire by whole-tree harvesting in jack pine forests: Effects on soil fertility and tree nutrition

Large areas of northern coniferous forests once naturally maintained by stand-replacing wildfires have shifted to an anthropogenic disturbance regime of clearcut harvesting followed by natural or artificial regeneration, with unknown consequences for soil biogeochemical processes. We used a comparative approach to investigate the effects of whole-tree harvesting (WTH) vs. stand-replacing wildfire (WF) on soil C and nutrient availability, and nutrition and growth of the succeeding stand, in jack pine (Pinus banksiana) forests of northern Lower Michigan. We compared total carbon (C), total nitrogen (N), potential N mineralization, and extractable phosphorus (P), potassium (K), calcium (Ca) and magnesium (Mg) among stands regenerated via WTH or WF in two age classes (4–7 years and 12–18 years). We also measured jack pine foliar nutrition and height growth in these same stands, as well as estimating the contribution of legacy dead wood to ecosystem nutrient capital in young stands. We found some evidence in support of our hypothesis that WTH would leave behind greater pools of soil C and N, but lower pools of P and base cations. However, the differences we observed were confined entirely to surface organic horizons, with the two disturbance regimes indistinguishable when viewed cumulatively to our maximum sampling depth of 30 cm. Estimates of nutrient pools in legacy wood inherited by young jack pine stands were also small in comparison to total soil pools (ranging from 1 to 9% depending on the element), suggesting that decomposition and nutrient release from this material is not likely to result in noticeable differences in soil fertility later in stand development. Similar levels of soil nutrients between WTH- and WF-origin stands were reflected in our measures of jack pine foliar nutrition and height growth, which were both unaffected by mode of stand origin. Results from this study suggest that soil nutrient levels following WTH fall within the natural range of variation produced by WF in these jack pine forests; however, comparison with a similar study on boreal jack pine suggests that latitudinal effects on O-horizon nutrient capital may influence the degree to which WTH matches the effects of WF on soil nutrient availability.     

Importance of weed control, fertilization, irrigation, and genetics in slash and loblolly pine early growth on poorly drained spodosols

Research reported here has evaluated interactions of factors limiting the biological potential of slash pine (Pinus elliottii var. elliottii Engelm.) and loblolly pine (P. Taeda L.). Water and nutrients were manipulated by irrigation, weed control, and intensive fertilization. Genetic factors were incorporated as (1) improved loblolly seedlings from a commercial clonal seed orchard and (2) slash seedlings from four fast-growing, open-pollinated seed-orchard clones. Three replications of a 2 × 2 × 2 factorial experiment were established on a Typic Haplaquod soil in northern Florida using four cultural treatments: no treatment (F0H0); weed control (F0H1); fertilizer (F1F0); and combination of weed control and fertilizer (F1H1) for each species. During the first year there were large, sustained responses to all cultural practices. Loblolly and slash pine showed similar growth responses and had a lengthened growing season (60–100 days). Slash pine growth was superior to that of loblolly. Growth responses observed during the first year continued. After 4 years, irrigation had no measurable effect on tree growth. Both pine species responded equally to F1H0 and F0H1, but slash retained a slight edge. Extra nutrients]available via fertilizer, reduced competition, or additional rooting space, increased slash-pine volume index from an average of 2.0 m³ ha⁻¹ (F0H0) to 11.90 and 13.59 m³ ha⁻¹ (F0H1 and F1H0, respectively). The F1H1 treatment produced the largest slash-pine response, diameter at breast height (D_bh) averaged 9.84 cm, height reached 5.02 m, and volume index rose to about 22.94 m³ ha⁻¹. With teh F1H1 treatment, average loblolly-pine height and volume exceeded that of the average slash pine. At age 4, the growth response due to increased nutrient supply was 13–29 times that of the best genetic response within F0H0 trees of the four slash-pine progenies. The best-growing slash-pine family (6–56) exceeded loblolly-pine growth under all four treatments and approximated the vigor shown by slash pine in the southern hemisphere.     

Clonal variation in crown structure, absorbed photosynthetically active radiation and growth of loblolly pine and slash pine

Crown structure, absorbed photosynthetically active radiation (APAR) and growth were analyzed in 300 replicated loblolly (Pinus taeda L.) and slash pine (Pinus elliottii Engelm. var. elliotti) clones to: (1) quantify genetic variation in crown structural traits, growth and APAR at the species, family and clonal levels; and (2) estimate within-family genetic and environmental influences on measured variables. Species and family-within-species differences were found in some growth traits, crown size, leaf area, APAR and branch angle. Loblolly pine developed larger crowns, exposed more leaf area with an acute angle, and intercepted more radiation than slash pine. Significant differences among clones within-family were found for stem volume and crown architecture. Loblolly pine and slash pine within-family, individual-tree broad-sense heritabilities ranged from 0.00 to 0.41 for growth and crown structural traits and most were between 0.10 and 0.25 when estimated from a combined analysis across families. Genetic correlations of crown size, leaf area and APAR with volume increment generally ranged from 0.60 to 0.75. This knowledge of the genetic interactions among growth and crown structural traits improves our understanding of how crown morphology affects light interception and stand development, and ultimately how these attributes can be incorporated in the selection of families or clones for the development of new crop tree ideotypes.     

Effects of silvicultural treatments and seasonal patterns on foliar nutrients in young post-fire Pinus halepensis forest stands

Tree growth and health status appear to be related to foliar nutrient contents. Foliar nutrient concentration might be the result of a complex interaction between soil nutrients and effective availability caused by climate, water and other site and treatment effects. This study examines foliar macronutrient (N, P, K, Ca, Mg) and organic C concentrations in Aleppo pine (Pinus halepensis) needles (between 5 and 18 months old), as well as time course variability (nine dates, from July 1999 to November 2001). Variability was assessed depending on quality site (two sites, Yeste and Calasparra; SE Spain) and seven silvicultural treatments including thinning, scrubbing, pruning and particular combinations of them. Foliar macronutrient concentrations for Aleppo pine in South-eastern Spain were slightly lower (N, P, K,) or higher (Mg, Ca) than the considered as adequate ranges for Aleppo pine and Pinus genera. However, our results agree well with other normal ranges reported for Aleppo pine in Spain and for other North American Pinus species such as P. elliottii, P. taeda and P. palustris. Site, treatment and date (season) affected significantly the foliar macronutrient and C concentration, although the most important was the date, likely due to the two growth periods per year that Aleppo pine has in Mediterranean sites. Silvicultural treatments affected foliar nutrient concentration, so that the concentrations of N, P and K were higher when treatments included thinning than those that did not. The contrary was true for Mg and Ca. However, treatments did not affect the time course of the concentration, i.e., seasonality was not broken due to treatments. Moreover, the effect of the treatments was markedly high along the first year after they were applied but the differences were attenuate 2 years later. Site affected the time course of N, K, Mg and C in a different way: while for N, K and C, at the end of study period, they were higher in Calasparra than in Yeste, for Mg the contrary was true. Nutrient ratios had a different behaviour regarding to single nutrient: although date was significant, we did not appreciate seasonality. In addition, some nutrient ratios were not affected by treatments (N/P, N/K, Ca/Mg,) or by site (N/Ca, K/Ca). Average foliar N concentration and Ca/Mg ratio explained significantly the mean diameter and height growth, so that higher is the foliar N concentration and lower is Ca/Mg, higher is the growth.