A method for quantifying changes in the epicuticular wax structure of Norway spruce needles

A method for quantifying changes in the epicuticular wax structure of spruce needles (Pieea abies) has been developed for the scanning electron microscope. This evaluation system, based on a five-point scale, classifies only the epistomatal waxes of current-year needles. Two criteria, the different crystal wax forms and the amount of changed wax as a percentage of the total stomatal area have been used for classification. Statistically significant results (accuracy below ± 10% at 5% error level) were obtained under the following conditions. Mixed needle samples were taken from two adjacent trees per site. These were in a dominant and predominant social position in the stand. The needles were collected from the 7^th whorl of sun crowns exposed to all wind directions. The wax quality was determined by evaluating three stomata per needle. One stoma was investigated at the needle base, one m the middle and the third at the apex of the needle. 24 needles were evaluated per site. SEM-Investigations of needle waxes of 25 Austrian forest sites with no local emission source (background sites) showed, that the mean wax quality ranged from 1.6 to 4.1, according to this classification method. Stepwise multiple regression analyses with site characteristics and element contents of the needles (nutrient elements, heavy metals) confirmed correlations between the wax quality and the manganese- and sulphur-content of the needles and also the average degree of crown defoliation.     

The effects of Arceuthobium sichuanense infection on needles and current‐year shoots of mature and young Qinghai spruce (Picea crassifolia) trees

Arceuthobium sichuanense is a hemiparasitic angiosperm that infects Qinghai spruce (Picea crassifolia Kom.) in Qinghai province, China, and causes severe damage to spruce forests in Qinghai‐Tibet Plateau. In this study, the impact of A. sichuanense infection on mature and young trees of Qinghai spruce was evaluated by examining needle and current‐year shoot morphology, needle water and nitrogen‐use efficiency (NUE) and needle nitrogen concentration. The most apparent effect of A. sichuanense infection was a significant reduction in both needle size distal to infection and current‐year shoot length in the infected branches (p < 0.001). Per cent reductions in needle and current‐year shoot length were similar between mature and young trees (58.9 vs. 56.3%; 59.7 vs. 62.9%). There was a high degree of correlation in foliar δ¹⁵N values between the dwarf mistletoe and its host trees (R² = 0.9017, p < 0.001), while the foliar δ¹³C values of A. sichuanense were similar to those of infected mature and young spruce trees. The dwarf mistletoe infection also resulted in a significant decrease in host needle N concentration and δ¹³C values (p < 0.001). The per cent reduction in needle N concentration in young trees was nearly twice as much as that in mature trees (20.49 vs. 11.54%), while the per cent reduction in needle δ¹³C values was similar between young and mature trees (?0.98 vs.?1.1‰). The NUE in mature trees was not affected by A. sichuanense infection, but the NUE in young trees was increased by the infection.     

Changes in shoot properties in relation to vertical positions within the crown of mature canopy trees of Abies mariesii and Abies veitchii

We investigated current shoot properties in two contrasting vertical positions (leader crown; LC, and lower branch; LB) within the crowns of mature trees of two subalpine conifer species, Abies mariesii and A. veitchii. For both LCs and LBs, shoot length decreased with increasing branching order. However, shoot properties were different between LCs and LBs. Shoots in LCs had more needle biomass per unit of shoot length. Shoots in sunny conditions pack needles closer along the shoot and intercept incoming light more completely. This causes the shoots in the LCs to have more needles. In contrast, less needle packing per unit shoot length in LBs results in the avoidance of mutual shading among needles in order to intercept limited light more effectively. Because branch systems in lower layers tend to be more shaded, the quantity of irradiance received by the shoots in LBs is smaller. Thus, reduced needle amounts on the shoots in LBs reflect the needle arrangement acclimating to the lower light availability. This study suggests the importance of changes in the properties of individual shoot as a component of a branch system and accordingly a whole-crown system in mature canopy trees of A. mariesii and A. veitchii.     

Photosynthesis,nutrient, growth and soil investigations of a declining Norway spruce (Picea abies) stand in the coastal region of Northern Germany

The research site, Wingst Compartment 123B, is a 68-year-old Norway spruce (Picea abies (Karst.)) stand located in the coastal area of northern Germany. This area receives high atmospheric inputs of ammonium and also has relatively high ozone concentrations (0.061 mg m⁻³).Ten trees were categorized as healthy to slightly damaged (3–29% needle loss) or severely damaged (49–71% needle loss). Apparent net photosynthetic rates were measured on detached branches at light saturation (1000 μE m⁻² s⁻¹). Needles were analyzed for chlorophyll, N, C, Ca, Mg, K, Zn, Mn and Fe.When compared to the healthy-to-slightly-damaged trees, the severely damaged trees tended to have higher rates of net apparent photosynthesis in the 1 and 2-year-old needles and similar rates in the current-year needles. All three needle ages from the severely damaged trees had higher average stomatal conductances to water vapor (g_s. Although the damaged trees had significantly less total chlorophyll in all needle ages sampled, there was no statistically significant difference in the chlorophyll a:b ratio between the healthy and severely damaged trees.Nitrogen contents of the current-year needles were slightly lower in the severely damaged trees. Carbon and calcium levels did not significantly differ between the damage classes, although the average Ca content of all younger needles was generally under the recommended sufficiency levels. The current-year needles of the severely damaged trees had significantly less magnesium (0.42–0.46 mg Mg g⁻¹) than those of the healthy trees, and all trees had Mg values in the deficiency range (< 0.7 mg g⁻¹). The severely damaged trees also had lower average potassium levels in the older needles.The annual volume increment per unit crown surface area declined with increasing crown damage. Trees with a 50% needle loss showed a 62% loss of volume increment.Soil investigations revealed conditions of high soil acidity and poor nutrient capacity. The low pH values (pH < 3.8) in 64% of all samples indicated a high risk of acid toxicity for plant roots in the investigated area.The significance of these results relative to the current ideas concerning forest decline is discussed.     

Needle nutrient status of Sirococcus shoot blight‐diseased and healthy Norway spruces

In this study the needle nutrient status of Sirococcus shoot blight‐diseased and healthy Norway spruces (Picea abies) was investigated. Element contents were determined in the current year and 3‐year‐old needles collected in December 2000 from 72 mature trees randomly distributed in the investigated stand. Half of the trees were severely affected by Sirococcus conigenus and had a reduced crown whereas the other trees were unaffected and vigorous. Compared with the uninfected trees, contents of Mg, Ca, P and Mn were significantly lower in the current year and 3‐year‐old needles of diseased trees. Furthermore, analyses of the nutrient contents of needles from severely affected trees revealed insufficient Mg and Ca supply and enhanced N/Mg and N/Ca ratios, when compared with threshold values. The results of this study suggest that imbalanced tree nutrition either has a significant impact on the expression of Sirococcus disease symptoms or fungal infection itself is affecting needle nutrient contents.     

Effects of Filtered Air and Misting Treatments on Cuticles of Red Spruce Needles on Whiteface Mountain,N.Y.

Two sets of large (8-20 m tall) red spruce (Picea rubens Sarg.) trees growing on Whiteface Mountain, N.Y., were used in branch chamber experiments. In the summer of 1988, the treatments were made on four healthy trees and consisted of: (1) open branches (N), and in chambers that were supplied with either: (2) ambient air (A); (3) charcoal-filtered air (T); or (4) charcoalfiltered air and deionized water misting when the forest was immersed in ambient cloud water (M). Each tree had all of the treatments. The treatments for the second year (summer of 1989) were made on four declining trees and consisted of: (1) open branches (NC) and chambers supplied with either (2) ambient air and mist (AA), (3) synthetic acid mist and ambient air (SA), (4) distilled water mist and ambient air (DA), (5) synthetic acid mist with filtered air (SF), and (6) distilled water mist with filtered air (DF). Foliar properties measurcd included: (1) microreflectivity, a measure of surface wax properties; (2) cuticle thickness; (3) cutinized cell wall thickness; (4) total cuticular layer thickness: (5) the mass of surface wax extracted from needle samples; and (6) contact angles, a measure of surface wetting. In addition, needles were collected and measured from healthy trees at lower and higher elevations. In the first experiment, reflectance from the wax in the stomata1 antechambers in treatment M was statistically significantly greater than all the other treatments. The cuticles of needles in treatment M were also statistically significantly thicker than the cuticles measured in all the other treatments (40% thicker than for the open branches). A plot of cuticular layer against elevation showed that the two filtered chambers (M and T) provided environments for cuticle development that mimic lower elevations. In 1989 the weather was cooler and more moist, and although the trees used in 1989 were declining, the results were similar to those of 1988. However, the additional treatments with synthetic acid mist resulted in reduced cuticle thickness. While the cuticles of the chamber trees growing at 1,170 m were, except for treatment M and T, generally thicker in 1989 than in 1988, they were still considerably thinner than those of spruce trees growing at the lower elevations. These results suggest regional scale air and precipitation quality have detrimental effects on red spruce needles at Whiteface Mountain.     

Epicuticular waxes,peroxidase activity and contents of chlorophyll,phaeophytin and carotenoids of needles from Pinus syivestris and picea abies as indicators of air pollution in the region of Stockholm

To trace any possible effects of air pollution stress and injuries on trees around a city centre, needles of Scots pine (Pinus Syivestris L.) and Norway spruce (Picea abies (L.) Karst.) from sites of different distances and directions from Stockholm were analysed. Comparisons were made between needles of different ages, a half to three and a half years old (C ‐ C + 3) for pine up to six and a half years old (C ‐ C + 6) for spruce. Epicuticular waxes, studied by scanning electron microscopy (SEM), showed age‐dependent but site‐independent changes. Different age‐patterns were found in the pine and spruce needles but no dust coverings or lesions were detected, irrespective of the sites. A proportionally increasing peroxidase activity (EC 1.11.1.7) with increasing needle‐age up to the third year was observed in both species. No certain indication of site‐effects, expressed by an unproportional increase of the enzyme activity with increasing needle‐age could be found. Chlorophyll a and b, phaeophytin a and b and carotenoid analyses on the corresponding materials only indicated disturbances in trees from some scattered sampling plots. Thus, neither could the pigment analyses reveal any signs of a general pollution stress in pine and spruce trees from the area in question.     

Within-crown structural variability of dwarfed mature Abies mariesii in snowy subalpine parkland in central Japan

We investigated mature dwarf Abies mariesii trees growing in conifer thicket–meadow parklands on a snowy subalpine plateau, where these dwarf trees are buried in the accumulated snow in winter. We focused on structural variation in the needles, shoots, and branchlets within different crown positions (leader crown vs lower crown) of the dwarf trees. In the leader crown, which appears above the snow surface earlier than the lower crown, current-year shoots and branchlets had greater total biomass, and foliage was more closely packed along the stem axis than in the lower crown, whereas current-year shoots in the leader crown had a lower needle mass ratio than in the lower crown. These results suggest that current-year shoots and branchlets in the leader crown have a specific structure that allows them to harvest more light, although construction and maintenance costs would be higher. In contrast, the structural characteristics of current-year shoots and branchlets in the lower crown efficiently concentrate incoming light by avoiding mutual shading within foliage, thus leading to increased biomass of photosynthetic needles within shoot and branchlet biomass. Such within-crown variability at various hierarchical levels from needles to branches in mature, but very dwarf, A. mariesii trees maintains the crown and allows survival within conifer clumps in areas of subalpine parklands that receive heavy snowfall.     

Leaf conductance of primary and mature leaves ofPinus roxburghii: A comparison

We used a null-balance porometer to measure leaf conductance of mature and primary leaves of natural seedlings, saplings, and trees ofPinus roxburghii (chir) during autumn at four sites at 1,320–1,930 m elevation in the central Himalaya of India. Our hypothesis that primary leaves had higher leaf conductance than mature leaves (needles in fascicles), based on measurements in other pines, was rejected. Comparisons on the same saplings and seedlings showed lower leaf conductance for primary leaves than for mature leaves. Primary leaves on seedlings and saplings also did not consistently have higher leaf conductance than mature leaves on near-by trees. Mature leaves on seedlings, however, did often have higher conductance than mature leaves on nearby trees. Mean values for leaf conductance (mmol m⁻² s⁻¹, based on total leaf surface area) ranged from 42–82 for mature leaves on trees, 60–121 for mature leaves on seedlings, and 28–67 for primary leaves on seedlings. Compared to published values for other pine species, conductance of mature leaves ofP. roxburghii is relatively low. Funding for this study was provided by grants from the United States National Science Foundation, INT-9312052 and INT-9404043. We appreciate the review of an early draft by Barbara Bond.     

Needle anatomy changes with increasing tree age in Douglas-fir

Morphological differences between old-growth trees and saplings of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) may extend to differences in needle anatomy. We used microscopy with image analysis to compare and quantify anatomical parameters in cross sections of previous-year needles of old-growth Douglas-fir trees and saplings at the Wind River Canopy Crane site in Washington and at three sites in the Cascade Mountains of Oregon. We also compared needle anatomy across a chronosequence of 10-, 20-, 40- and 450-year-old Douglas-fir trees from the Wind River site. Anatomy differed significantly between needles of old-growth trees and saplings at all sites, suggesting a developmental change in needle anatomy with increasing tree age. Compared with needles of old-growth trees, needles of saplings were longer and had proportionately smaller vascular cylinders, larger resin canals and few hypodermal cells. Astrosclereids, which sequester lignin in their secondary cell walls and occupy space otherwise filled by photosynthetic cells, were scarce in needles of saplings but abundant in needles of old-growth trees. Needles of old-growth trees had an average of 11% less photosynthetic mesophyll area than needles of saplings. The percentage of non-photosynthetic area in needles increased significantly with increasing tree age from the chronosequence of 10-, 20-, 40- and 450-year-old trees at the Wind River site. This reduction in photosynthetic area may contribute to decreased growth rates in old trees.     

Mycobiota in needles of Abies alba with and without symptoms of Herpotrichia needle browning

The ascomycete Nematostoma parasiticum (syn.: Herpotrichia parasitica) is commonly perceived as the causal agent of the so‐called Herpotrichia needle browning in silver fir (Abies alba). However, its fruitbodies are rarely present on symptomatic needles, which are also colonized by many presumably saprotrophic fungi. We compared the internal colonization of healthy and symptomatic needles on two sites in Poland. In addition, the endophytic mycobiota in needles of various age was recorded on two other sites without disease symptoms. Fungi were isolated from 95.6% of the dead needles and from 62.9% of the living needles on symptomatic trees, whereas on healthy trees, only 45.0% of the needles were colonized internally. Colonization frequency increased with needle age. From a total of 2017 isolates, 116 fungal taxa were identified. Frequency of many species was influenced by needle type. Anthostomella formosa, Gloeosporidiella sp., Hypoxylon fragiforme, Xylaria hypoxylon and X. polymorpha were the most common fungi isolated from living needles. In symptomless living needles, fungi occurred significantly more often in the basal than in the apical parts. In dead needles, the most common fungi were Alternaria alternata, Paraconiothyrium sporulosum, Fusarium sp., Mollisia cinerea, Rhizoctonia sp., Rhizosphaera oudemansii, Thysanophora penicillioides, Xylaria hypoxylon and X. polymorpha. Rhizoctonia sp. was the most frequently isolated fungus in dead needles (23.4%) but occurred rarely also in living needles (0.3–1.1%). The supposed pathogen N. parasiticum was detected only sporadically (at most in 0.6% of the needles). Our findings demonstrate the need for understanding the role of Rhizoctonia sp. in Herpotrichia needle browning disease aetiology.     

Pine needle consumption and bark stripping by sheep grazing annual pastures in young stands of widely spaced Pinus radiata and P. pinaster

The consumption of the pine needles and the stripping of bark by sheep grazing annual pastures in three open stands of P. radiata and one of P. pinaster were measured. The P. radiata were four years old and the densities were 250, 500 and 750 trees ha^-1. The P. pinaster included two, three and four year-old trees and the density was 440 trees ha^–1.Needles were eaten immediately after the sheep were admitted and this continued throughout the year. Within ten weeks of the start of winter grazing, sheep stocked at 7–10 ha^–1 had eaten about half of the accessible needles on both species where densities were less than500 trees ha^–1. The estimated consumption was about 260 g sheep^–1 day^–1 for the P. radiata. Bark stripping then commenced and continued throughout spring and summer on most plots. In some plots at low tree density, bark was stripped from more than 35% of the trees. In denser stands, needle consumption per tree was less and bark was stripped from less than 5% of the trees.Bark damage was most common on the smaller trees, and severity of stripping was negatively correlated with tree size. Of all the trees with bark damage, 22% of the P. pinaster and 14% of the P. radiata had more than half of the stem circumference stripped. A number of these will probably suffer both stem distortion and growth retardation, and some may die.Methods of reducing the incidence of bark damage are discussed.     

Impacts of Swiss needle cast on overstory Douglas-fir forests of the western Oregon Coast Range

Tree-ring analysis was applied to assess the impacts of the fungal disease Swiss needle cast on the radial growth of mature Douglas-fir (Pseudotsuga menziesii) forests in the western Oregon Coast Range. Although considered endemic to the Pacific Northwest, Swiss needle cast has significantly lowered productivity in Douglas-fir forests only in the past 20–30 years. To date, studies on Swiss needle cast impacts have almost exclusively involved young (<30 years) plantation trees. To better describe the history of Swiss needle cast and its impacts on older (>80 years) trees, we extracted tree cores from dominant and codominant Douglas-fir and western hemlock (Tsuga heterophylla) in three even-aged stands in western Oregon. In the least affected stand growth rates of both species did not significantly differ, while at the most severely diseased site Douglas-fir radial growth was reduced by as much as 85%. Growth reductions likely associated with Swiss needle cast were dated to as early as 1950, though the most severe impacts occurred after 1984. An index of Swiss needle cast severity significantly (p < 0.01) related to instrumental records of air temperatures such that warm conditions from March through August were associated with reduced radial growth at the most severely affected site. Overall, this study demonstrates that even mature forests of natural origin are susceptible to severe growth reductions by Swiss needle cast, that warmer spring and summer temperatures are associated with Swiss needle cast impacts, and that the disease appears to be increasing in severity.     

Defoliation and growth relationships for mid-rotation Sitka spruce attacked by the green spruce aphid, Elatobium abietinum (Walker) (Homoptera: Aphididae)

Green spruce aphid (Elatobium abietinum) feeds primarily on the 1-year-old and older needle leaves of spruce (Picea spp.) and is a major defoliator of commercial Sitka spruce plantations in the UK and other maritime regions of north-west Europe. The impact of E. abietinum on mid-rotation (23-28-year-old) Sitka spruce in Radnor Forest, in mid-Wales, was determined by comparing aphid population densities, defoliation and growth between plots of trees that were either treated with insecticide or were left untreated. The experimental treatments were maintained for 5 years and over this period (2002-2006) peak E. abietinum densities in the untreated plots varied from 5 to 36 aphids per 100 needles. These densities, which were representative of low to moderate rates of infestation, were associated with low rates of defoliation (0-8%), but they had a significant (P ? 0.01) impact on mean radial increment (RI) and mean volume increment (VI). In 2005, the year with the highest aphid populations, peak densities in the untreated plots averaged 14 aphids per 100 needles and this rate of infestation reduced RI by 17%, VI by 10% and the dry weight of current-year needles by 10%. On average, across all years, infestation by E. abietinum reduced mean annual VI by 6%. Comparisons with previous studies on the impact of the aphid on 4-year-old, 7-year-old and 9-year-old Sitka spruce (Straw et al., 2005) indicate that defoliation by E. abietinum, on a per capita basis, decreases as trees grow older, but that the impact on VI increases. The different growth response of young and mid-rotation Sitka spruce to infestation is related to differences in canopy structure, particularly in the ratio of current-year needles to older needles, and the greater demands on photosynthetic production in older trees that arise from the need to support an increasing quantity of non-photosynthetic structural tissues in branches, stem and roots.     

The relationship between the needle age and the growth rate in Scots pine (Pinus sylvestris): a retrospective analysis by needle trace method (NTM)

Variation in annual radial and height increments (RI and HI, respectively) as well as the number of needles belonging to different needle sets of successive annual shoots, determined by the needle trace method (NTM), was studied retrospectively in Scots pine stands, one in southern Finland and two in Estonia. The connection between different needle sets and tree growth was studied by correlation and covariance analyses. The numbers of needles of the three youngest needle sets were positively correlated with growth, whereas the number of needles older than 3 years had no significant correlation. The covariance analysis emphasized the negative relationship of older needle sets with tree growth. It is generally concluded that the two or three youngest needle sets are important and all older needles minimally contribute to growth, possibly because of their small quantity. Older needles may remain on the tree if assimilation is impeded for some reason, e.g. due to premature loss in the two youngest needle sets. The trees having attached old needles grew less than those that had already totally lost these needle sets.     

Differential effects of plant ontogeny and damage type on phloem and foliage monoterpenes in jack pine (Pinus banksiana)

Coniferous trees have both constitutive and inducible defences that deter or kill herbivores and pathogens. We investigated constitutive and induced monoterpene responses of jack pine (Pinus banksiana Lamb.) to a number of damage types: a fungal associate of the mountain pine beetle (Dendroctonus ponderosae Hopkins), Grosmannia clavigera (Robinson-Jeffrey & R.W. Davidson); two phytohormones, methyl jasmonate (MJ) and methyl salicylate (MS); simulated herbivory; and mechanical wounding. We only included the fungal, MJ and mechanical wounding treatments in the field experiments while all treatments were part of the greenhouse studies. We focused on both constitutive and induced responses between juvenile and mature jack pine trees and differences in defences between phloem and needles. We found that phytohormone applications and fungal inoculation resulted in the greatest increase in monoterpenes in both juvenile and mature trees. Additionally, damage types differentially affected the proportions of individual monoterpenes: MJ-treated mature trees had higher myrcene and β-pinene than fungal-inoculated mature trees, while needles of juveniles inoculated with the fungus contained higher limonene than MJ- or MS-treated juveniles. Although the constitutive monoterpenes were higher in the phloem of juveniles than mature jack pine trees, the phloem of mature trees had a much higher magnitude of induction. Further, induced monoterpene concentrations in juveniles were higher in phloem than in needles. There was no difference in monoterpene concentration between phytohormone applications and G. clavigera inoculation in mature trees, while in juvenile trees MJ was different from both G. clavigera and simulated herbivory in needle monoterpenes, but there was no difference between phytohormone applications and simulated herbivory in the phloem.     

Effect of soil chemical properties on growth,foliation and nutrition of Norway spruce stand affected by yellowing in the Bohemian Forest Mts., Czech Republic

In 1998, a fertilizer experiment aiming to investigate the effects of slow-release N, P, K and Mg fertilizer (SILVAMIX Mg NPK^®) on a 60-year-old spruce stand with symptoms of yellowing was established. In this paper, trees were selected to investigate the relation between annual diameter increment, yellowing, foliation, needle and soil chemical properties: ten from the fertilized treatment (F), ten green trees from the control (CG) and ten yellow trees from the control (CY). CG and CY trees were growing in close proximity at a distance of only several meters apart under the same soil conditions. In treatment F, increased annual diameter increment, improved foliation, needle Mg concentration, plant-available Mg and P concentrations in the soil and absence of yellow trees were recorded 7 years after a single application of the fertilizer. During the last 15 years, annual growth increment and foliation of CY trees have continuously decreased while relatively stable values were recorded for CG trees and increased for F trees. In 2006, CG and CY trees differed significantly in Mg concentration in needles, foliation, yellowing and annual diameter increment. Although differences in soil chemical properties between CG and CY treatments were not significant, lower concentrations of plant-available Mg²⁺ and higher concentrations of H⁺ and Al³⁺ were found in soils under CY trees. There was a negative correlation between soil concentration of Mg and yellowing, but this correlation was relatively weak, indicating that there is no simple relation between soil and needle concentrations of Mg. In the investigated locality, the “new type” of yellow tree decline has been a long-term gradual process.     

马尾松二代无性系生长和针叶N/P化学计量特征

[目的]在土壤有效P匮乏的立地条件下,揭示针叶N、P含量及其化学计量比与马尾松生长性状的相关性。[方法]以马尾松二代育种群体内30个无性系为试验对象,野外调查11年生时马尾松二代无性系树高、胸径等生长性状与针叶N、P含量及其化学计量比的相关性。[结果]结果表明:(1)各生长性状在马尾松二代无性系间差异显著,且无性系重复力较高,均达到0.95以上。马尾松成熟针叶和当年生针叶N、P含量及N/P在二代无性系间差异均达显著水平;(2)相关分析表明,马尾松二代无性系树高、胸径与成熟针叶和当年生针叶N含量、N/P均呈显著或极显著正相关,与成熟针叶N/P的相关系数分别为0.608 6和0.542 8,与当年生针叶N/P的相关系数分别为0.515 9和0.679 2;(3)基于主要生长性状、针叶N和P含量及其化学计量比,可将30个马尾松二代无性系聚为4类:类群1无性系树高、胸径等生长表现优良,成熟针叶N/P较高,而当年生针叶N/P相对较低;类群2无性系树高、胸径等生长表现较好,成熟针叶和当年生针叶N/P值在30个无性系中居中间水平;类群3占试验无性系总数的一半,树高、胸径等生长表现一般,成熟针叶与当年生针叶N/P相差较大,大部分无性系成熟针叶N/P较低,而当年生针叶N/P相对较高;类群4树高、胸径等生长表现较差,成熟针叶和当年生针叶N/P均较低。[结论]在贫P立地下,马尾松二代亲本无性系C13(7750×1126)和C14(3201×1123)各生长指标均较高,可能是通过调整当年生针叶和成熟针叶的养分分配,增加了当年生针叶对限制其生长的P素的再利用和再吸收,促进了N素的吸收,调整了当年生针叶和成熟针叶的N/P,以形成特定的养分分配机制。     

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.     

Effect of dothistroma needle blight on needle and shoot lengths

The needles of pine trees are indicative of the overall health of the tree, and their length is affected by many factors. This study describes the effect of high levels of pathogen infection on lengths of both needles and shoots. Dothistroma septosporum is an important foliar pathogen of pines causing necrosis and premature defoliation with successive years of high infection leading to growth reduction and in extreme cases tree death. Corsican pine trees with all foliage infected by D. septosporum had needles of primary, secondary and tertiary shoots reduced by 31.8%, 44.0% and 64.2%, respectively, compared to non‐infected trees. Needle lengths were reduced both in the upper and lower portions of the canopy, with a greater reduction lower in the canopy. Trees with high levels of infection had shorter shoots in the upper canopy with fewer, shorter needles on these shoots compared to trees with low levels of infection. The results demonstrate the substantial negative effect on needle and shoot lengths of trees with high levels of D. septosporum infection, comparable to factors such as water and nutrient availability known to have a strong influence on these parameters. The reductions in length reduce the photosynthetic capacity of the tree and compound the immediate reductions caused by necrosis and premature defoliation. These effects on needle and shoot lengths contribute to the reductions in volume growth of affected trees and, moreover, are longer lasting than the immediate effects of necrosis and premature defoliation.