Driving factors for natural tree rejuvenation in anthropogenic pine (Pinus sylvestris L.) forests of NE Germany

The rejuvenation ecology of three main tree species in anthropogenic pine (Pinus sylvestris L.) forests is explored in our study. We focus on the scale of micro-plots, which provide the safe sites for tree rejuvenation. We thrive on the multi-factorial relationship of tree establishment and driving ecological factors using a large dataset from pine stands in NE Germany and applying multivariate analyses. The success of the establishment of the investigated focal tree species Fagus sylvatica L., Quercus petraea Liebl. and Pinus sylvestris L. is, on general, mostly affected by three factors, i.e. water balance of the upper soil layers, browsing pressure, and diaspore sources. Our investigations on the micro-plot scale revealed species-specific differences. For beech saplings <50 cm growth height, primarily the availability of water, indicated by available water capacity (AWC), thickness, quality, and structure of the organic layer, silt and humus content in the topsoil, and the lack of a dense competitive herb layer, were identified as most important factors. On the contrary, oak seems hardly be restricted by hydrologic and/or trophic deficits in the topsoil or humus layer. In conclusion and comparison to Fagus sylvatica L., we assume for Quercus petraea Liebl. advantages in natural regeneration processes under sub-continental climate conditions and thus under the scenarios of climate change. Pinus sylvestris L. regeneration in our investigation area occurs only in a narrow niche. We conclude with regard to future forest development and the objective of stand conversion with low management intensity that oak should be favoured within natural stand regeneration.     

Regeneration of Rocky Mountain bristlecone pine (Pinus aristata) and limber pine (Pinus flexilis) three decades after stand-replacing fires

Rocky Mountain bristlecone pine (Pinus aristata) and limber pine (Pinus flexilis) are important high-elevation pines of the southern Rockies that are forecast to decline due to the recent spread of white pine blister rust (Cronartium ribicola) into this region. Proactive management strategies to promote the evolution of rust resistance and maintain ecosystem function require an improved understanding of the role of disturbance on the population dynamics of both species and environmental conditions that favor seedling establishment. We examined patterns of bristlecone and limber pine regeneration across the perimeters of three, 29-year-old, high-severity burns in northern, central, and southern Colorado: Ouzel, Badger Mountain, and Maes Creek, respectively. Both species exhibited a very protracted regeneration response to these fires. Bristlecone pine regeneration was concentrated near burn edges and beneath surviving seed sources. This spatial pattern is consistent with limitations incurred by wind-dispersal, also borne out by the low occurrence of seedling clusters. Relative to unburned stands, the absolute abundance of bristlecone pine generally increased only on plots retaining some surviving trees. Limber pine regeneration pattern varied between sites: high in the burn interior at Ouzel, concentrated at burn edges at Badger, and mostly in unburned stands at Maes. Clark’s Nutcracker dispersal of limber pine in each study area was indicated by high seedling distance from possible seed sources and high frequencies of clustered stems. Except at Ouzel, the absolute abundance of limber pine decreased in burns. Across sites, establishment by both species was boosted by nearby nurse objects (rocks, fallen logs, and standing tree trunks), a relationship that extended out at least as far as the closest three such objects, usually found within 50 cm. Fire decreased the frequency of Pedicularis but increased Castilleja and Ribes species (alternate hosts of white pine blister rust), though only one species, R. cereum, was positively associated with either pine species. We conclude that regeneration of bristlecone and limber pine may benefit from natural disturbance or proactive management creating appropriately sized openings and microtopographic structure (e.g., abundant fallen logs); however, beneficial responses may require many decades to be achieved.     

黑松和马尾松苗受松材线虫侵染后部分化学物质的响应

为研究松材线虫侵染对寄主植物生理生化物质代谢的影响,以黑松和马尾松4 5年生苗为实验材料,测定了接种松材线虫对2种寄主植物的营养物质和次生代谢物质含量的变化及其规律。结果显示:接种松材线虫初期,黑松和马尾松的总糖含量均较高,随接种时间的延长,总糖含量呈不断下降趋势;黑松的可溶性糖含量一直降低并明显低于对照,马尾松的可溶性糖含量在侵染前期与对照相比变化不明显,而15 d后快速降低;黑松和马尾松的可溶性蛋白含量侵染前期均低于对照,后升高,再降低;黑松的单宁含量明显高于对照,马尾松的单宁含量接种第1天略低于对照,第3天后开始升高;黑松和马尾松的总酚含量均在侵染前期高于对照,而后降低。这些物质的变化趋势显示松材线虫侵染不同寄主植物的生理反应。     

Endophyte-mediated resistance against white pine blister rust in Pinus monticola

Induced resistance responses, including fungal endophyte-mediated resistance, have been well studied in both agricultural crops and grass systems. Yet, the effect of these processes and symbionts in forest trees is poorly known. Fungal endophytes have been found in all conifer forest systems examined to date and have been hypothesised to be involved in resistance-mediated responses. However, in the absence of functional studies the influence of these endophytes on the extended phenotype of the host plant is unclear. In this study we demonstrate that fungal endophytes from Pinus monticola were effective at increasing survival in host plants against the exotic pathogen Cronartium ribicola, which is responsible for the devastating disease called white pine blister rust. Seedlings previously inoculated with fungal endophytes lived longer than endophyte-free seedlings and also showed some reduction in white pine blister rust disease severity. This endophyte-mediated resistance was found to be effective over time, indicating persistence, and is hypothesised to be a form of induced resistance. Overall, this suggests fungal endophytes may play a determinative role in the structure of biological communities and could provide a useful alternative or ancillary management tool for combating pests and diseases.     

Vulnerability of Pinus cembra L. in the Alps and the Carpathian mountains under present and future climates

Proactive management should be applied within a forest conservation context to prevent extinction or degradation of those forest ecosystems that we suspect will be affected by global warming in the next century. The aim of this study is to estimate the vulnerability under climate change of a localized and endemic tree species Pinus cembra that occurs in the alpine timberline. We used the Random Forest ensemble classifier and available bioclimatic and ecological data to model present and future suitable areas for P. cembra and estimate its current and future vulnerability. Future projections for years 2020, 2050 and 2080 were simulated using two IPCC Special Report on Emission Scenarios run under four global climate models.     

Influence of woody and herbaceous competition on microclimate and growth of eastern white pine (Pinus strobus L.) seedlings planted in a central Ontario clearcut

The influence of woody and herbaceous plant competition, either alone or in combination, on microclimate and growth of planted eastern white pine (Pinus strobus L.) seedlings was examined over four consecutive growing seasons in a central Ontario clearcut. Treatments that manipulated the comparative abundance of these two plant functional groups significantly affected light availability, soil moisture, and air and soil temperature regimes. These microclimate alterations, coupled with the relative competitiveness of herbaceous and woody vegetation, corresponded to temporal changes in vegetation cover and dominance. The more rapid colonization and growth of the herbaceous plant community, dominated by bracken fern (Pteridium aquilinum) and ericaceous shrubs (Kalmia sp., Vaccinium sp.), resulted in this form of vegetation being a comparatively important early competitor for soil moisture. As the woody plant community, dominated by naturally regenerated trembling aspen (Populus tremuloides Michx.), grew in height and leaf area, it became a comparatively strong competitor for both light and soil moisture. For all vegetation treatments combined, white pine seedling growth responses were strongly correlated with total cover of competing vegetation and its relative influence on above- and belowground microclimatic variables. Higher total cover of competing vegetation was generally associated with lower light and soil moisture availability and cooler soil temperatures. Multiple regression analyses indicated that pine seedling relative height growth increased with soil moisture content and growing season soil heat sum, while seedling relative diameter and relative volume growth increased with light availability.     

马尾松钙调素基因的克隆及响应松材线虫侵染的表达特征

松材线虫病是由松材线虫(Bursaphelenchus xylophilus)侵染引起的严重病害,引起重大的经济和生态损失。钙调素(CaM)是真核生物中Ca2+主要传导蛋白,本研究采用RT-PCR方法,首次从马尾松中克隆获得CaM,命名为pmCaM,并检测了其响应松材线虫侵染的表达特征。序列分析表明:pmCaM完整的开放阅读框核苷酸序列为450bp,编码149个氨基酸的蛋白质;该蛋白含有4个EF-hand结构域,具有钙调素的典型特征,与其它植物的CaM蛋白均有较高同源性。实时荧光定量PCR分析显示:接种松材线虫后30 180 min时间段,马尾松苗根、茎、叶器官中的pmCaM均下调表达,但不同器官间显著下调表达的时间点存在差异;同时,不同器官pmCaM表达量变化均存在特异的随时间发展的波动特征,其中发现,根茎pmCaM在松材线虫接种45 min时均处于表达量高峰。本研究结果显示pmCaM表达响应了松材线虫侵染,揭示pmCaM可能参与了调控松材线虫-马尾松互作早期的钙信号响应。     

Population biology of sugar pine (Pinus lambertiana Dougl.) with reference to historical disturbances in the Lake Tahoe Basin: Implications for restoration

Historical logging, fire suppression, and an invasive pathogen, Cronartium ribicola, the cause of white pine blister rust (WPBR), are assumed to have dramatically affected sugar pine (Pinus lambertiana) populations in the Lake Tahoe Basin. We examined population- and genetic-level consequences of these disturbances within 10 sugar pine populations by assessing current population structure and trends for 1129 individuals, genetic diversity for 250 individuals, and frequency of WPBR-resistance for 102 families. Logging had occurred in 9 of 10 sites and fire suppression was evident in all stands. High density of white fir (Abies concolor) is often an indicator of fire suppression and we found a negative relationship between sugar pine survivorship and white fir basal area (r² = 0.31). C. ribicola was present in 90% of stands (incidence range: 0-48%) and we found a significant relationship between mean host survivorship and disease incidence (r² = 0.46). We estimated population growth rates (λ) from size-based transition matrices. For six of 10 sugar pine populations λ was ?1.0, indicating that these populations appear to be stable; for four populations, λ was <1.0, indicating populations that may be in decline. A population specific drift parameter, c_i, which is a measure of genetic differentiation in allele frequencies relative to an ancestral population, ranged from 0.009 to 0.048. Higher values of c_i indicate greater genetic drift, possibly due to a bottleneck caused by historical logging, other agents of mortality or much older events affecting population sizes. Effects of drift are known to be greater in small populations and we found a negative relationship between sugar pine density and c_i (r² = 0.36). Allele frequency of the Cr1 gene, responsible for WPBR-resistance in sugar pine, averaged 0.068 for all populations sampled; no WPBR infection was found in one population in which the Cr1 frequency was 0.112. Historical disturbances and their interactions have likely influenced the population biology of sugar pine in the Tahoe Basin; for some populations this has meant reduced population size, higher genetic drift, and poor survival of small- and intermediate-sized individuals. Possible management strategies include restoring population numbers, deploying WPBR-resistance, treating stands to promote natural sugar pine regeneration, and enhancing genetic diversity.     

Effect of patch size and isolation on mating patterns and seed production in an urban population of Chinese pine (Pinus tabulaeformis Carr.)

Fragmentation is a critical issue for tree populations because the creation of small patches can reduce local population size and increase isolation, both of which can promote inbreeding and its negative consequences, as well as loss of genetic diversity. To test the hypothesis that patch size and isolation influence mating patterns or seed production in forest trees, we utilize the spatial array of trees of an urban population of Chinese Pine (Pinus tabulaeformis Carr.) planted in patches around Beijing. Our design includes 28 urban patches, with patch size ranging from 1 to 2000 adult trees and isolation (edge distance index) ranging from 37.5 m to 245.8 m. We examined the average number of seeds per cone and percentage of viable seed per cone for each patch as measures of seed production. By utilizing seven paternally inherited chloroplast microsatellite loci, we estimated the mating pattern parameters for each patch, including the level of selfing, the amount of immigrant pollen and the effective number of pollen sources (N_ep). Using a general linear model selection procedure based on AIC value, we found patch size was the best predictor of the selfing and immigration rate; smaller patches had a higher selfing and immigration rate. Small patches with one adult had relatively high N_ep which indicates connectivity among urban patches. However, due to the reduced amount of immigrant pollen and limited diversity of local pollen, intermediate sized patches (with 5–10 adults) had the lowest N_ep among the study patches. For patches with more than 10 adults, N_ep was increased with patch size. The percentage of viable seeds per cone significantly decreased with patch size, indicating a possible negative consequence of inbreeding. The effect of patch size on mating patterns and seed production suggests that the patches of trees experience less connectivity than trees within continuous forest. These findings indicate that forest management practices should emphasize the maintenance of an optimal patch size because, despite the fact that tree species show the potential for long distance pollen movement, the number of local trees strongly influences the mating patterns.     

云南松天然林及人工林群落结构和物种多样性比较

通过对云南省滇中地区云南松天然林和人工林的群落结构调查及物种多样性研究,结果表明:干扰较少的云南松天然林具有较为稳定、合理的群落结构,其乔木层优势种群具有较强的自然更新能力和长远的演替潜力,并且物种多样性和生境多样性。干扰较少的云南松天然林的这些特性均优于干扰较大的云南松天然林,干扰较大的云南松天然林又优于人工林。随着森林起源过程中干扰(人为干扰和自然干扰)程度的逐渐增加,云南松群落的稳定性、生物多样性、协调平衡能力及自然修复能力相应下降;而入侵种紫茎泽兰的分布范围和数量则相应上升,提示其或可作为森林群落生物多样性、干扰程度和森林健康评价的指示生物。     

Restoring longleaf pine (Pinus palustris Mill.) in loblolly pine (Pinus taeda L.) stands: Effects of restoration treatments on natural loblolly pine regeneration

Historical land use and management practices in the southeastern United States have resulted in the dominance of loblolly pine (Pinus taeda L.) on many upland sites that historically were occupied by longleaf pine (Pinus palustris Mill.). There is currently much interest in restoring high quality longleaf pine habitats to such areas, but managers may also desire the retention of some existing canopy trees to meet current conservation objectives. However, fast-growing natural loblolly pine regeneration may threaten the success of artificially regenerated longleaf pine seedlings. We evaluated the establishment and growth of natural loblolly pine regeneration following different levels of timber harvest using single-tree selection (Control (uncut, residual basal area ∼16 m²/ha), MedBA (residual basal area of ∼9 m²/ha), LowBA (residual basal area of ∼6 m²/ha), and Clearcut (complete canopy removal)) and to different positions within canopy gaps (approximately 2800 m²) created by patch cutting at two ecologically distinct sites within the longleaf pine range: Fort Benning, GA in the Middle Coastal Plain and Camp Lejeune, NC in the Lower Coastal Plain. The density of loblolly pine seedlings was much higher at Camp Lejeune than at Fort Benning at the end of the first growing season after harvesting. Following two growing seasons, there were no significant effects of canopy density or gap position on the density of loblolly pine seedlings at either site, but loblolly pine seedlings were taller on treatments with greater canopy removal. Prescribed fires applied following the second growing season killed 70.6% of loblolly pine seedlings at Fort Benning and 64.3% of seedlings at Camp Lejeune. Loblolly pine seedlings were generally less than 2 m tall, and completeness of the prescribed burns appeared more important for determining seedling survival than seedling size. Silvicultural treatments that include canopy removal, such as patch cutting or clearcuts, will increase loblolly pine seedling growth and shorten the window of opportunity for control with prescribed fire. Therefore, application of prescribed fire every 2-3 years will be critical for control of loblolly pine regeneration during restoration of longleaf pine in existing loblolly pine stands.     

马尾松人工林地力维护研究进展

针对地力衰退与维护这一林业研究热点,文中从林分生产力、土壤理化性质、土壤养分、土壤微生物、土壤酶活性、自毒作用6个方面阐述了马尾松人工林的地力现状。众多研究表明,马尾松人工林尚未出现地力衰退,其产生地力衰退的原因主要是不合理的营林措施。同时从轮作、混交林、发展林下植被、林地施肥、生态系统管理等方面介绍了马尾松人工林地力维护进展,并对马尾松人工林地力维护研究趋势进行了展望。     

马尾松谷胱甘肽过氧化物酶PmGPX6基因cDNA克隆及转化拟南芥耐旱性初步研究

[目的]克隆马尾松谷胱甘肽过氧化物酶基因,并对其进行功能研究。[方法]采用RACE技术克隆基因c DNA序列,实时荧光定量PCR检测基因在马尾松干旱胁迫下的表达模式,花序浸泡法转化拟南芥,并对转基因与野生型拟南芥的生长表型和根系生长进行分析。利用荧光显微镜技术对转基因拟南芥不定根进行GFP荧光检测。[结果]克隆到1个871 bp的GPX基因全长c DNA序列,命名为Pm GPX6。Pm GPX6包括513 bp的完整开放阅读框,编码170个氨基酸残基。Pm GPX6蛋白与油松Pt GPX蛋白同源性达95%。Pm GPX6在马尾松根中高表达,茎、叶中表达量低。在干旱胁迫下,Pm GPX6在根、茎、叶中的表达量均在第15天达到最大,随后出现下降趋势。过表达Pm GPX6与野生型拟南芥植株在正常水分条件下表型与根长差异不大,但在干旱胁迫下,转基因植株根系更长。转基因拟南芥根在蓝色光激发下能发出强烈的绿色荧光,表明Pm GPX6基因能高效表达。[结论]推测Pm GPX6可能参与马尾松干旱胁迫应答。     

Oleoresin flow and chemical composition of Corsican pine (Pinus nigra subsp. laricio) in response to prescribed burnings

Oleoresin is an important defensive made up of acidic and neutral diterpenes together with a variable fraction of monoterpenes and sesquiterpenes. It plays a major role in plant-herbivore defense. Fire, or prescribed burning, can be a stress factor leading to metabolic changes in pine needles. Prescribed burning is used to: (i) restore fire to the ecosystem, (ii) recreate natural disturbance dynamics and (iii) reduce fuel loadings and fire risks. To our knowledge, no study has been performed on the effects of fire on the oleoresin defense system in pine trees, apart from the measurement of resin flow. We examined the effect on oleoresin production (resin flow) and the chemical composition (volatile and diterpene components) in Corsican pine (Pinus nigra subsp. laricio (Poir.) Maire var. corsicana (Loud.) Hyl.). This species is an ecologically and economically important pine in France. The chemical composition of samples was studied using HS-SPME GC and GC/MS for the volatile fraction and ¹³C NMR for diterpenes. A considerable increase in resin production and, consequently, an increase predominantly in diterpenes together with volatile compounds were observed at stations which had sustained recent prescribed burnings. Resin flow in trees in low-intensity prescribed burning after 4 years were no different than trees in unburned plots. Resin flow can be considered as a response to thermal stress. The major components of the two fractions were α-pinene (66.9–75.6%) and levopimaric acid (19.8–23.0%), respectively. The chemical composition of the resin remained unchanged after prescribed burning, even though resin production increased. No insect attacks were observed after low-intensity prescribed burning of P. laricio.     

Multiscale spatial variation of the bark beetle Ips sexdentatus damage in a pine plantation forest (Landes de Gascogne,Southwestern France)

Bark beetles are notorious pests of natural and planted forests causing extensive damage. These insects depend on dead or weakened trees but can switch to healthy trees during an outbreak as mass-attacks allow the beetle to overwhelm tree defences. Climatic events like windstorms are known to favour bark beetle outbreaks because they create a large number of breeding sites, i.e., weakened trees and for this reason, windthrown timber is generally preventively harvested and removed. In December 1999, the southwest of France was struck by a devastating windstorm that felled more that 27 million m³of timber. This event offered the opportunity to study large-scale spatial pattern of trees attacked by the bark beetle Ips sexdentatus and its relationship with the spatial location of pine logs that were temporally stored in piles along stand edges during the post-storm process of fallen tree removal. The study was undertaken in a pure maritime pine forest of 1300 ha in 2001 and 2002. We developed a landscape approach based on a GIS and a complete inventory of attacked trees. During this study more than 70% of the investigated stands had at least one tree attacked by I. sexdentatus  . Spatial aggregation prevailed in stands with n≥15$n\ge 15$ attacked trees. Patches of attacked trees were identified using a kernel estimation procedure coupled with randomization tests. Attacked trees formed patches of 500–700 m² on average which displayed a clumped spatial distribution. Log piles stemming from the sanitation removals were mainly distributed along the large access roads and showed an aggregated spatial pattern as well. The spatial relationship between patches of attacked trees and log pile storage areas was analyzed by means of the Ripley’s statistic that revealed a strong association at the scale of the studied forest. Our results indicated that bark beetle attacks were facilitated in the vicinity of areas where pine logs were stored. The spatial extent of this relationship was >1000 m. Similar results were obtained in 2001 and 2002 despite differences in the number and spatial distribution of attacked trees. The presence of a strong “facilitation effect” suggests that log piles should be removed quickly in order to prevent outbreaks of bark beetles.     

欧洲黑杨PnEXPA1基因多态性与稳定碳同位素比率的关联分析

对欧洲黑杨α-expansin基因PnEXPA1的SNP多态性与水分利用效率相关性状稳定碳同位素比率(δ13C)进行了关联分析。利用SNaPshot技术对PnEXPA1基因内11个SNP位点进行了基因型分型,发现各SNP位点优势基因型均为纯合,且其频率高于杂合基因型。关联分析显示:SNP8和SNP12在采用单因素方差分析(ANOVA)和一般线性模型(GLM)2种方法时均与δ13C值显著关联。SNP8为exon 1内的无义突变,可解释6.620%的表型变异;而SNP12位于intron 1中,遗传贡献率为6.613%;这2个SNP位点与SNP9、SNP13共同位于一个高连锁不平衡(LD)的单倍型块中。SNP8与SNP12的TT基因型无性系均具有较高的δ13C值,为优势基因型。     

Susceptibility of ponderosa pine, Pinus ponderosa (Dougl. ex Laws.), to mountain pine beetle, Dendroctonus ponderosae Hopkins,attack in uneven-aged stands in the Black Hills of South Dakota and Wyoming USA

Mountain pine beetle, Dendroctonus ponderosae Hopkins can cause extensive tree mortality in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests in the Black Hills of South Dakota and Wyoming. Most studies that have examined stand susceptibility to mountain pine beetle have been conducted in even-aged stands. Land managers increasingly practice uneven-aged management. We established 84 clusters of four plots, one where bark beetle-caused mortality was present and three uninfested plots. For all plot trees we recorded species, tree diameter, and crown position and for ponderosa pine whether they were killed or infested by mountain pine beetle. Elevation, slope, and aspect were also recorded. We used classification trees to model the likelihood of bark beetle attack based on plot and site variables. The probability of individual tree attack within the infested plots was estimated using logistic regression. Basal area of ponderosa pine in trees ≥25.4 cm in diameter at breast height (dbh) and ponderosa pine stand density index were correlated with mountain pine beetle attack. Regression trees and linear regression indicated that the amount of observed tree mortality was associated with initial ponderosa pine basal area and ponderosa pine stand density index. Infested stands had higher total and ponderosa pine basal area, total and ponderosa pine stand density index, and ponderosa pine basal area in trees ≥25.4 cm dbh. The probability of individual tree attack within infested plots was positively correlated with tree diameter with ponderosa pine stand density index modifying the relationship. A tree of a given size was more likely to be attacked in a denser stand. We conclude that stands with higher ponderosa pine basal area in trees >25.4 cm and ponderosa pine stand density index are correlated with an increased likelihood of mountain pine beetle bark beetle attack. Information form this study will help forest managers in the identification of uneven-aged stands with a higher likelihood of bark beetle attack and expected levels of tree mortality.     

芽梢斑螟危害马尾松雄球花的初步研究

马尾松(Pinus massoniana Lamb.)是我国分布最广、数量最多的松树,是组成亚热带地区大面积森林的主要针叶树种,也是淮河以南酸性土荒山造林的先锋树种[1-2]。马尾松花粉有良好的医疗和保健价值。汉代《神农本草经》记载松花粉气味甘温无毒,主治心腹寒热邪气,利小便,消淤血,久服轻身益     

Growth and nutrition of Pinus radiata in response to fertilizer applied after thinning and interaction with defoliation associated with Essigella californica

Infestations of Essigella californica following the installation of post-thinning fertilizer trials in Pinus radiata plantations provided an opportunity to examine the impact of repeated defoliation over a period of 8 years (1997–2005). Replicated treatments (n = 4) of nil fertilizer (control), N (300 kg ha⁻¹) as urea, P (80 kg ha⁻¹) and S (45 kg ha⁻¹) as superphosphates were applied immediately after thinning at three sites and this was followed by a second application of NPS fertilizers 6 years later with N applied at 300 kg ha⁻¹ as urea and ammonium sulphate and P at 80 or 120 kg ha⁻¹. Defoliation of untreated P. radiata gradually increased to 50% over a period of 8 years. Basal area growth was negatively correlated with average defoliation for two consecutive post-fertilizer periods of 6 and 2 years. Growth responses to fertilizer varied considerably between sites but the largest improvement in growth was due to NPS fertilizer, this increased basal area by 30–80%. Application of N fertilizer raised total N levels in foliage and increased defoliation with a commensurate loss in growth under conditions of deficiencies of S or P. Repeated infestations gradually increased the percentage of trees with severe defoliation (>80% loss of foliage) indicating that nutrient-deficient trees have a reduced capacity for foliage recovery between episodes of peak infestation. In contrast, treatment with N fertilizer in combination with S- and P-corrected deficiencies of these nutrients, raised levels of total N in foliage and reduced defoliation to approximately 20%. Basal area growth responses to NPS fertilizers reflected improved nutrition as well as reduced insect damage. The reduction in defoliation under conditions of balanced tree nutrition was most likely due to enhanced needle retention following correction of P deficiency as well as greater availability of nutrients enabling a more vigorous recovery of P. radiata after an episode of E. californica activity. Treatment with fertilizer therefore reduced the long-term impact of aphid damage and improved growth of P. radiata.