The productivity of pine plantations in relation to previous land use

Summary Differences in the productivity of establishedP. radiata plantations on pasture and forest soils were found to be reproducible withP. radiata seedlings in a glasshouse environment. The growth of seedlings on pasture soil exceeded that of seedlings on native forest soil (the ‘primary pasture effect’). After a history of pine on both native and pasture soils a residual effect of pasture on seedling growth was evident (the ‘secondary pasture effect’). However, the effect of a history of pine plantation (the ‘pine effect’) was to decrease the productivity of both native and pasture soil as assessed by seedling growth. These effects were not related to changes due to land management in mycorrhizal infection or in soil structure. The analysis of seedling growth leads to the conclusion that soil fertility, particularly the availability of nitrogen and phosphorus, has changed.     

The chemical composition of pine foliage in relation to the population dynamics of the pine beauty moth,Panolis flammea,in Scotland

Summary This paper reports part of a study to determine why damaging outbreaks of the pine beauty moth, Panolis flammea (D & S) (Lepidoptera: Noctuidae), in Scotland are frequent on lodgepole pine but do not occur on Scots pine, and why outbreaks on lodgepole pine are mainly confined to trees growing in deep unflushed peat. The elongation of shoots and the growth of needles of Scots pine occurred later in the season than did those of lodgepole pine. The foliage of Scots pine generally had a higher level of nitrogen, and consistently had a higher level of phosporus, but had a consistently lower level of tannins than that of lodgepole pine during the period when the larvae were feeding each year. The nitrogen content of the foliage of lodgepole pine growing in an iron pan soil was generally higher than that of lodgepole pine growing in deep peat during the same period but there were no general differences in the phosphorus or tannin contents of lodgepole pine in the 2 soil types. These findings suggest that Scots pine is a more suitable host plant than lodgepole pine and that the foliage of lodgepole pine growing in deep peat is not more suitable than lodgepole pine growing in an iron pan soil. On the basis of the chemical analyses used in this study, it is concluded that the abundance of pine beauty moth in Scotland is not strongly influenced by the nutritional suitability of its host plants.     

Exotic pine plantations and indigenous Lepidoptera in Australia

Temperate regions of Australia have extensive commercial plantations of Monterey pine, Pinus radiata D. Don. Replacement of indigenous forests by P. radiata is likely to have significant effects on assemblages of native Lepidoptera, and has been considered a major threat to native fauna through displacing native species. However, many species of Lepidoptera, including ennomine geometrid moths in the genus Chlenias Guenèe, have adopted P. radiata as a larval host. Comparisons were made of oviposition preferences and nutritional ecology of Chlenias auctaria Guenée on P. radiata and two native host plants [Acacia mearnsii De Wild. (Mimosaceae) and Eucalyptus obliqua LHérit (Myrtaceae)]. Females showed no significant oviposition preference for any of the three hosts. Growth of sixth instar larvae was significantly less on P. radiata than on the native hosts. Pupal weights were significantly lower, suggesting that the fitness of moths reared on P. radiata will be significantly reduced. The life history strategy of C. auctaria, which includes dispersal of first instar larvae by ballooning, may predispose this species to feed on poor quality hosts, and this may be common to other polyphagous species found feeding on P. radiata in southern hemisphere plantations. The impact of P. radiata may have a long term effect on lepidopteran communities, beyond the simple replacement of indigenous host plants leading to extirpations of feeding specialists.     

Role of phosphorus and nitrogen in photosynthetic and whole plant carbon gain and nutrient use efficiency in eastern white pine

Summary In white pine (Pinus strobus) seedlings grown in five forest soils from New York State, net photosynthetic capacity (Amax) plant^-1 was correlated with total foliar N plant^-1 (r ²=0.57), but was more highly correlated with total foliar P plant^-1 (r ²=0.82). There was no relationship (r ²<0.01) between Amax [g leaf]^-1 and foliar N [g leaf]^-1 for the pooled data set, but there was a significant (P<0.001), but weak (r ²=0.20) positive relationship between Amax [g leaf]^-1 and foliar P [g leaf]^-1 across all soils. However, within two of the five soils leaf N concentration was a significant (P<0.05) determinant of photosynthetic capacity. Due to differences in soil nutrient availabilities a large range in foliar P:N ratio (0.02–0.15) was observed, and the proportion of leaf P:N appeared to control Amax [g leaf N]^-1. Whole plant nitrogen (NUE) and phosphorus (PUE) use efficiencies were well correlated with whole plant P:N ratio. In addition, NUE was well correlated with Amax [g leaf N]^-1 and PUE was well correlated with Amax [g leaf P]^-1. However, NUE was not well correlated with PUE, and Amax [g leaf N]^-1 was not well correlated with Amax [g leaf P]^-1. These results indicated that P and/or N limitations were important components of photosynthetic nutrient relations in white pine grown in these five soils and suggest that both P and N and their proportions should be considered in analyses of photosynthesis-nutrient relations.     

Root development and configuration in intensively managed radiata pine plantations

Summary In south-east Australia, where radiata pine (Pinus radiata D. Don) is grown on sandy soils low in nutrients and short of water, early establishment, and rapid growth to canopy closure lead to increased productivity. At this stage demands for nutrients and water are high, and trees respond vigorously to silvicultural inputs.For several months after transplanting in winter roots are confined within a narrow planting wedge, low temperature restricts new root growth and slows recovery from water stress in plants. From spring, depending upon the configuration and vigour of the roots transplanted, lateral roots extend radially throughout the soil.Although there were small decreases in concentration of roots radially from the stems of very young trees, such spatial differences disappeared between ages 2 and 3, so that rooting density was independent of distance from the stem. The pattern of vertical distribution of lateral roots was not influenced by age and 80–90% of the lateral roots were within the top 30 cm soil. Roots developed rapidly as the trees grew towards canopy closure, but in general the rooting densities of these pines are among the lowest reported for plants. In rapidly growing trees approaching canopy closure, the secondary thickening of the lateral roots was sufficient to double the weight of roots without altering root length.Knowledge about root growth and root configuration during the early phase of plantation development will assist management decisions where intensive silviculture is practiced, and hence ensure the most efficient use of nutrients and water.     

Nutrient proportions in foliage of semi-mature loblolly pine

Summary Nitrogen and phosphorus limitations to growth are common in many loblolly pine (Pinus taeda) stands. Interactions of these nutrients may complicate interpretation of foliar nutrient analysis for predicting response to forest fertilization. Proportions of foliar nutrient concentrations (and the changes in these proportions following fertilization) were examined in 36 semi-mature loblolly pine plantations in the southeastern United States. Mean proportions of nutrient concentrations (NPKCaMg) for non-fertilized stands were 1009.336.517.29.2. Potassium and phosphorus were higher. Nitrogen fertilization generally decreased the PN ratio and enhanced growth, indicating a nitrogen deficiency in most stands under study. Additions of nitrogen and phosphorus together yielded a significant increase in the PN ratio. Effects of fertilization effects on other nutrient concentration ratios were also examined.Paper No. 9401 of the Journal Series of the North Carolina Agricultural Research Service, Raleigh, NC 27695.     

Stem deformity inPinus radiata plantations in south-eastern Australia:

Visual symptoms of stem deformity similar to those of Cu deficiency are common inP. radiata established on fertile sites previously used for agricultural production in south-eastern Australia. In this study, Cu fertiliser was applied at rates of 0, 2, 5, 10, 20 and 50 kg ha⁻¹ to deformedP. radiata at ages 3 and 6 years. Available soil Cu and contents of Cu in the foliage increased significantly only in the younger plantation. Cu fertiliser did not affect growth nor did it improve stem form. Levels of N, Cu and Cu/N ratios in foliage of straight and deformed trees were similar. However, contents of Cu in apical buds were significantly lower in deformed trees. It was concluded that stem deformity inP. radiata as observed on these fertile pasture sites, cannot be corrected by application of Cu fertiliser. Differences in Cu levels in apical buds of straight and deformed trees suggest that Cu may still be involved in this syndrome. There was also no indication of other nutrient deficiencies that could be associated with the deformity.     

Inferring relatedness and heritability using molecular markers in radiata pine

A set of eight unlinked microsatellite markers was used to estimate relatedness among 355 individuals of a Pinus radiata breeding population. The average performance of open-pollinated progeny of each individual, for wood density, was considered to represent the phenotype of all 355 individuals. Marker-based estimates of relationship were compared with the pedigree-based coefficients of relationships. The phenotypic similarity among all pairs of individuals was regressed on marker-estimated relatedness to estimate the inheritance of wood density. The marker-based estimate of heritability was compared with that obtained using classical quantitative genetic methods. Overall, a low correlation (0.13) was observed between marker-based and pedigree-based estimates of relatedness. After discarding negative estimates of relatedness, the average coefficient of relationship among known groups of maternal half-sibs, full-sibs and unrelated individuals, increased from 0.24 to 0.29 (0.25 expected), from 0.43 to 0.48 (0.50 expected) and from –0.04 to 0.15 (0 expected), respectively. Marker-based and conventional estimates of heritability of wood density were 0.79 and 0.38, respectively. However, by using only marker loci with expected Hardy–Weinberg frequencies, marker-based estimate of heritability was 0.33, which is very similar to that obtained from conventional approaches. The use of molecular markers to understand quantitative genetic variation is discussed.     

Identification of an agar constituent responsible for hydric control in micropropagation of radiata pine

Tissue cultured Pinus radiata grown on media containing agar as the gelling agent display toxic symptoms and poor long-term shoot survival, however it does have the attribute of hydric control, through a mechanism which, until now, has not been elucidated. Gelrite as an alternative gelling agent is clearly non-toxic but results in hyperhydric (vitrified) tissues. In an effort to overcome these problems, the controlling mechanism found in agar was examined. Hydric control was shown to be effected by a non-gelling, cold-water soluble constituent of a commercial agar, rather than by physical properties of the gel. It could be separated from low molecular weight components of the agar responsible for the toxic symptoms by dialysis. It was identified as being an agaroid-type xylogalactan bearing pyruvate and sulphate substituents. Improved management of gelling agents in culture medium has contributed substantially to a thirty fold increase in propagation rates.     

Evidence for recombination and segregation of virulence to pine in a hybrid cross between Gibberella circinata and G. subglutinans

Two species associated with the Gibberella fujikuroi species complex, G. circinata (the cause of pitch canker in pines) and G. subglutinans (avirulent on pine), were found to have limited interfertility in hybrid crosses. MAT idiomorphs, polymorphisms in the histone H3 gene, vegetative compatibility, and virulence phenotypes were used to verify recombination. The MAT idiomorphs appeared to be assorting independently, but the histone H3 haplotype was disproportionately represented by that of the G. subglutinans parent. Ninety-eight percent (45/46) of the progeny tested were vegetatively incompatible with both parents. All F₁ progeny were avirulent to pine, but a wide range of virulence was restored through a backcross to the virulent parent (G. circinata). Attempts at hybrid crosses using other isolate combinations were rarely successful (1/26). This limited interfertility supports retention of G. circinata and G. subglutinans as separate species, but offers opportunities to characterize the inheritance of virulence to pine.     

13C NMR study of pine needle decomposition

The quality of substrates in plantation forest litter, and their chemistry, can influence decomposition and N cycling. We studied the decomposition of Pinus radiata D. Don needles suspended on branches in windrows, for 3 yr after clear-cutting, using improved solid-state ¹³C NMR and chemical analysis. The NMR spectra suggested that the concentration of condensed tannins was 12–22%, and showed they were chemically altered during the period 4–12 months after clear-cutting. The spectra showed no evidence for further chemical modification of the tannins during the second or third years. Data for P. radiata needle decomposition in New Zealand indicated rapid loss of mass in the first 3 months, and condensed tannins did not appear to prevent mineralization of C or N. The tannin and lignin concentrations increased with decomposition of the needles, which was consistent with the early mineralization of readily available C compounds.     

Nitrogen nutrition,photosynthesis and carbon allocation in ectomycorrhizal pine

Summary Studies examined net photosynthesis (Pn) and dry matter production of mycorrhizal and nonmycorrhizalPinus taeda at 6 intervals over a 10-month period. Pn rates of mycorrhizal plants were consistently greater than nonmycorrhizal plants, and at 10 months were 2.1-fold greater. Partitioning of current photosynthate was examined by pulse-labelling with¹⁴CO₂ at each of the six time intervals. Mycorrhizal plants assimilated more¹⁴CO₂, allocated a greater percentage of assimilated¹⁴C to the root systems, and lost a greater percentage of¹⁴C by root respiration than did nonmycorrhizal plants. At 10 months, the quantity of¹⁴CO₂ respired by roots per unit root weight was 3.6-fold greater by mycorrhizal than nonmycorrhizal plants. Although the stimulation of photosynthesis and translocation of current photosynthate to the root system by mycorrhiza formation was consistent with the source-sink concept of sink demand, foliar N and P concentrations were also greater in mycorrhizal plants.Further studies examined Pn and dry matter production ofPinus contorta in response to various combinations of N fertilization (3, 62, 248 ppm), irradiance and mycorrhizal fungi inoculation. At 16 weeks of age, 6 weeks following inoculation with eitherPisolithus tinctorius orSuillus granulatus, Pn rates and biomass were significantly greater in mycorrhizal than nonmycorrhizal plants. Mycorrhizal plants had significantly greater foliar %P, but not %N, than did nonmycorrhizal plants. Fertilization with 62 ppm N resulted in greater mycorrhiza formation than either 3 or 248 ppm. Increased irradiance resulted in increased mycorrhiza formation.     

樟子松人工固沙林稳定性的研究

提出了人工林稳定性的内涵及其评价标准．分析了影响沙地樟子松人工林稳定性的各种干扰因子,通过对沙地樟子松人工林造林成活、生长情况、对不良外界环境的抗御能力、林分寿命、林分结构和生产力的综合评价分析,认为沙地樟子松引种人工林基本上是稳定的;而生长在立地较好的丘间低地的林分,其稳定性大于沙丘顶部的林分;从抗枯梢病角度看,幼林的抗性大于中林;从林分密度和生产力角度来看．红花尔基沙地樟子松林稳定性高于章古台．     

Long-term needle litterfall of a Scots pine Pinus sylvestris stand: relation to temperature factors

Summary Needle litterfall of a Scots pine was caught over 24 years (1962–1986) with litter-traps in a Scots pine stand in southeastern Finland. The age of the trees averaged 111 years in 1962. The stand was naturally recruited and only minor silvicultural treatments occurred during its history. Litterfall showed great year-to-year variation, the minimum being 18 g/m² (in 1968) and maximum 213 g/m² (in 1973). There was no overall trend in the amount of litterfall, and the age of the stand was thus not important in determining the needle fall. We used time domain time series analysis (ARIMA) and standard climatic data (temperature, precipitation) to investigate the relationship of litterfall to climatic factors. Mean July temperature was clearly correlated with needle litterfall. High temperature in July coincided with enhanced litterfall in the same and the next year. Litterfall enhanced litterfall in the same and the next year. Litterfall increased also after high temperatures during March–April, but only in the same year. In addition to these the litterfall had a 4-year self-dependency. This is approximately the same as the mean longevity of needles in the study area. Altogether the time series model we propose covers about 90% of the variance of the original time series.     

Transformation and regeneration of loblolly pine: shoot apex inoculation with Agrobacterium

Loblolly pine (Pinus taeda L.) is the mostimportant tree species in US commerce and has much to gain through geneticengineering. This species can be transformed using particle bombardment andAgrobacterium; however, the regeneration of plants fromtransgenic tissues has been difficult and the recovery of transgenic plants hasbeen rare. A shoot-based and genotype-independent transformation methodemploying Agrobacterium tumefaciens was used to facilitaterecovery of plants and permit the transformation of elite germplasm. Shootsfrom4–6 week old seedlings and adventitious shoots from culture wereinoculated with A. tumefaciens EHA101 (pGUS3), or EHA105(pSSLa.3), subjected to selection and regenerated. Shoots that survivedexhibited expression of the uidA gene (GUS) in a patterncharacteristic of the either the CaMV35S promoter (pGUS3), or the larch RbcSpromoter (pSSLa.3) transferred. Recovered plants were screened using PCRamplification. Southern DNA analyses and amplification of the T-DNA borderjunction confirmed genomic integration of both transferreduidA and nptII genes. In this proofofconcept study, the overall recovery of P. taeda shoots wasfair (10–20%), while recovery of intact rooted plants was poor (>1%)due to difficulty in rooting. Recovery of intact rooted plants from inoculatedshoots of P. eldarica and P. radiatawas more efficient (10–30%). The addition of a shoot multiplication stepand effective rooting protocols will improve the efficiency of this genotypeindependent transformation method in P. taeda, and inotherPinus spp.     

Short-term effects of thinning and liming on forest soils of pitch pine and Japanese larch plantations in central Korea

The influences of thinning (50% of standing density) and liming (Ca+Mg, 2 Mg ha⁻¹) on soil chemical properties were investigated for 2 years (2001, 2002) in 40-year-old pitch pine (Pinus rigida Mill.) and 44-year-old Japanese larch (Larix leptolepis Gord.) plantations established on similar soils. In general, soil properties varied significantly among plantations and treatments. For both plantations, thinning significantly increased soil organic C (SOC) concentrations whereas there were no significant changes in soil pH and Ca and Mg concentrations. In addition, thinning increased total soil N and Na concentrations for the pitch pine plantation and available P concentration for the Japanese larch plantation in the second year after the treatment. Liming did not affect soil chemical characteristics for the pitch pine plantation except for Na concentration. However, for the Japanese larch plantation, liming significantly increased soil pH and K, Ca and Mg concentrations and decreased SOC and total soil N concentrations. For both plantations, soil Al concentration did not change after thinning and liming and decreased exponentially with increased pH values. The increases in SOC and total soil N concentrations after thinning were possibly due to increases in decomposition of organic matter and root death. Although differences were not statistically significant, soil available P concentration tended to increase at early stages of liming for both plantations. These results suggested that thinning and liming seemed to regulate soil chemical properties for pitch pine and Japanese larch plantations established on similar soils.     

Non-morphogenic metabolism during shoot induction in radiata pine cotyledon explants

Summary The effects of excision, light and cytokinin (N⁶-benzyladenine) on¹⁴C-acetate metabolism in cotyledons ofPinus radiata (D. Don) were determined.¹⁴CO₂ was released and the distribution of radioactivity into lipids, sugars, organic acids and amino acids was determined. While light and cytokinin generally caused some increase in metabolism, the effect of excision, i.e., wounding, was most pronounced. Specific metabolites examined (citrate, malate, succinate, alanine, aspartate, glutamate and glutamine) were at least 50% greater in¹⁴C-labeling in excised cotyledons as compared to intact seedlings. This enhancement of wound metabolism would mask possible morphogenically-related changes occurring at that time. This research was supported by the Natural Sciences and Engineering Research Council of Canada Grant A-6467 to T.A. Thorpe.     

Pine plantations in south eastern Australia support highly impoverished ant assemblages (Hymenoptera: Formicidae)

Pitfall trap surveys of ants were undertaken in mature and intermediate-aged Pinus radiata plantations and parental eucalypt forest in central Victoria, Australia. Ant assemblages of pine plantations are an impoverished subset of the eucalypt forest assemblages. Very few ant morphospecies, most of them in very low abundance, were found in unthinned intermediate-aged pines, and the numbers were substantially higher in older, thinned pines in which understorey vegetation is considerably more developed. Implications for plantation management and conservation of epigaeic ant assemblages are discussed.     

The change of soil carbon stocks and fine root dynamics after land use change from a native pasture to a pine plantation

A published meta-analysis of worldwide data showed soil carbon decreasing following land use change from pasture to conifer plantation. A paired site (a native pasture with Themeda triandra dominant, and an adjacent Pinus radiata plantation planted onto the pasture 16 years ago) was set up as a case study to assess the soil carbon reduction and the possible reason for the reduction under pine, including the change in fine root (diameter <2 mm) dynamics (production and mortality). Soil analysis confirmed that soil carbon and nitrogen stocks to 100 cm under the plantation were significantly less than under the pasture by 20 and 15%, respectively. A 36% greater mass of fine root was found in the soil under the pasture than under the plantation and the length of fine root was about nine times greater in the pasture. Much less fine root length was produced and roots died more slowly under the plantation than under the pasture based on observations of fine root dynamics in minirhizotrons. The annual inputs of fine root litter to the top 100 cm soil, estimated from soil coring and minirhizotron observations, were 6.3 Mg dry matter ha⁻¹ year⁻¹ (containing 2.7 Mg C and 38.9 kg N) under the plantation, and 9.7 Mg ha⁻¹ year⁻¹ (containing 3.6 Mg C and 81.4 kg N) under the pasture. The reduced amount of carbon, following afforestation of the pasture, in each depth-layer of the soil profile correlated with the lower length of dead fine roots in the layer under the plantation compared with the pasture. This correlation was consistent with the hypothesis that the soil carbon reduction after land use change from pasture to conifer plantation might be related to change of fine root dynamics, at least in part.     

Testing for linkage disequilibrium in the New Zealand radiata pine breeding population

Linkage analysis is commonly used to find marker-trait associations within the full-sib families of forest tree and other species. Study of marker-trait associations at the population level is termed linkage-disequilibrium (LD) mapping. A female-tester design comprising 200 full-sib families generated by crossing 40 pollen parents with five female parents was used to assess the relationship between the marker-allele frequency classes obtained from parental genotypes at SSR marker loci and the full-sib family performance (average predicted breeding value of two parents) in radiata pine (Pinus radiata D. Don). For alleles (at a marker locus) that showed significant association, the copy number of that allele in the parents was significantly correlated, either positively or negatively, with the full-sib family performance for various economic traits. Regression of parental breeding value on its genotype at marker loci revealed that most of the markers that showed significant association with full-sib family performance were not significantly associated with the parental breeding values. This suggests that over-representation of the female parents in our sample of 200 full-sib families could have biased the process of detecting marker-trait associations. The evidence for the existence of marker-trait LD in the population studied is rather weak and would require further testing. The exact test for genotypic disequilibrium between pairs of linked or unlinked marker loci revealed non-significant LD. Observed genotypic frequencies at several marker loci were significantly different from the expected Hardy-Weinberg equilibrium. The possibilities of utilising marker-trait associations for early selection, among-family selection and selecting parents for the next generation of breeding are also discussed.