Photosynthesis in Norway spruce seedlings infected by the needle rust Chrysomyxa rhododendri

Chrysomyxa rhododendri (DC.) De Bary is a needle rust with a host shift between Rhododendron sp. and Norway spruce (Picea abies (L.) Karst.), penetrating only the new developing flushes of the conifer. Because little is known about its effects on trees, we investigated several parameters related to photosynthesis in artificially infected 3-year-old Norway spruce seedlings. The potential efficiency of photosystem II (PSII; derived from chlorophyll fluorescence measurements) was reduced in infected current-year needles as soon as disease symptoms were visible, about three weeks after inoculation. Two weeks later, photosynthetic O(2) evolution (P(max)) of infected needles was less than 20% of control needles, whereas respiratory O(2) uptake (R(D)) was about three times higher than that of control needles. Nonstructural carbohydrate concentrations were about 60% of control values in all parts of the shoots of infected trees. Photosynthetic inhibition was associated with marked decreases in chlorophyll concentration and chlorophyll a/b ratio but only a small reduction in carotenoid concentration. In infected trees, P(max) of noninfected 1-year-old and 2-year-old needles was 50 and 80% higher than in the corresponding age class of needles of control trees. Estimation of potential daily net dry mass production, based on P(max), R(D), specific leaf area, carbon content and needle biomass, indicated that seedlings infected once were able to produce 60%, and those infected twice only 25%, of the dry mass of controls. We conclude that afforestation and regeneration of Norway spruce is seriously impaired in regions where seedlings are frequently attacked by Chrysomyxa.     

Dry mass production in seedlings of Norway spruce infected by the needle rust Chrysomyxa rhododendri

The aim of this study was to assess the losses in productivity of Norway spruce (Picea abies) due to the attack by Chrysomyxa rhododendri, a rust that only penetrates new developing flushes. Three-year-old seedlings were artificially infected once or twice in two consecutive years and the dry mass was determined at the end of the growing season. Dry mass accumulation of all parts including roots was significantly reduced if more than about one-third of the young foliage was infected. Shoot and root dry mass of once severely infected seedlings were 32 and 48% less than that of controls. This deficit could not be compensated during a following rust-free year. When seedlings were infected in two consecutive years, dry mass accumulation was reduced by 58% compared with controls. Results indicate that rejuvenation of and afforestation with Norway spruce can seriously be impaired by Chrysomyxa at the alpine timberline where Rhododendron sp. (the telial host of the parasite) is widespread.     

Temporal variation in airborne spore concentration of Chrysomyxa rhododendri: correlation with weather conditions and consequences for Norway spruce infection

The fungal pathogen Chrysomyxa rhododendri undergoes a host shift between Rhododendron spp. and Picea abies and has a considerable negative impact on the latter by infecting the new sprouting needles and reducing photosynthesis, growth and seedling survival. Repeated high infection rates in the Central European Alps were reported in recent years, and although the life cycle of the pathogen is well understood, knowledge on temporal patterns of host infection, spore dispersal and influence of weather conditions is limited. We analysed the period of needle infection by a staggered application of fungicide on twigs of Norway spruce. Seasonal and diurnal patterns of spore dispersal were investigated using a Burkard volumetric spore trap, and airborne spore concentrations were correlated with several weather parameters. Needle infection occurred within 3 weeks (19 June–9 July). Basidiospores (infecting Norway spruce) were trapped from May to July, and airborne spore concentration was positively correlated with air temperature, global radiation and wind speed, and negatively with air humidity and precipitation. Aeciospores (infecting rhododendrons) appeared from August to October, and high concentrations were significantly associated with rain events. Needle infection started with bud sprouting and was limited either by decreasing spore concentrations or morphological and chemical differentiation of the needles, which prevented infection. Both spore types showed distinct periods of dispersal based on the life cycle of the pathogen, and the variability in concentration within these periods was explained by local weather conditions. The correlation analysis indicated different mechanisms of spore dispersal for both spore types and, with respect to global warming, more favourable conditions for the pathogen in future. The results can explain the high year‐ and site‐specific variation in Chrysomyxa infection intensities of P. abies and may be helpful for developing successful strategies to control the pathogen and protect alpine spruce forests.     

Stand development of Norway spruce dominated subalpine forests of the Swiss Alps

In densely populated regions, forests can help protect communities and infrastructures from natural hazards such as avalanches and rockfall. To promote the protective function, substantial efforts are made to actively manage forest stands. In 2009 alone the Swiss government invested more than 60 million sfr for the maintenance of protection forests. However, to date there has been no comprehensive evaluation of how the structural development of actively managed stands differs from that of passively managed stands in the Alps. Over the past century the structure of Norway-spruce dominated subalpine forests of the Swiss Alps has been changing and it is not clear how these changes affect the potential protective function of these forests, as well as other forest functions such as wildlife habitat. Furthermore, it is not clear how stand dynamics and structural changes differ between stands that are actively managed and those that are passively managed, and thus to what degree active efforts of forest management are contributing to stands that actually have a greater protective function than passively managed forests.In this study, we analyzed the dynamics of subalpine forests using time series of forest inventory data and examined the influence of active vs. passive management, exposition, distance to treeline, elevation and slope steepness on stand structure and dynamics. We analyzed data from 395 plots in dense, subalpine, spruce-dominated forests in the Swiss Alps. Data were collected during three distinct periods (1983-1985, 1993-1995 and 2004-2006) as part of the Swiss National Forest Inventory (NFI). Using regression trees and multivariate statistics, we investigated which factors have the most important influence on tree growth and stand development.Overall, forest density increased significantly over the last 20 years and the predominance of dense forests increased at higher elevations. However, forest density has not increased in actively managed forests over the past 10 years. In passively managed stands, density was higher on south-facing slopes than on north-facing slopes. The volume and density of dead wood has increased over the last 20 years in both actively managed and passively managed forests. Active management over the last 20 years has maintained forest conditions that adequately maintain stands’ protection functions in the Swiss Alps. However, stand development, especially increasing density, in the passively managed stands of the Swiss Alps suggests that the majority of passively managed stands also provide adequate protective functions against rock and avalanche hazards without the high costs of active management.     

Co‐occurrence of the ascomycete Lophodermium piceae and the rust fungus Chrysomyxa abietis in Norway spruce needles

The frequency of needles and the proportion of needle segments infected by Lophodermium piceae were compared in symptomless and Chrysomyxa abietis‐infected, 1‐year‐old needles of Picea abies. In late spring, symptomless needles from both rust‐infected and healthy saplings were sampled. In addition, rust‐infected, totally chlorotic needles and needles with chlorosis along about half their length from the diseased trees were examined. In all three stands, the proportion of segments infected by L. piceae was larger in the rust‐infected half of the needle than in the symptomless half; but the difference was statistically significant in only one of the stands. The proportion of L. piceae‐infected segments among the nonrust‐infected needles was the same as that found for the uninfected half of rust‐infected needles (after correction for size differences). No differences in the proportion of L. piceae‐infected segments were found between the totally chlorotic, rust‐infected needles and the green needles of diseased or healthy trees.     

Effects of repeated fertilization in young Norway spruce forests

Several studies have shown the positive effect of nitrogen fertilization on conifer growth. In young Norway spruce (Picea abies) stands, an additional effect of including a mixture of other nutrients has often, but not always, been found. We studied effects of repeated fertilization in 28 stands with young Norway spruce in central Norway. The treatments consisted of plots without nutrient addition (Control), fertilization with 150?kg?N?ha^?1 (150?N), and fertilization with 150?kg?N plus addition of P, K, Mg, B, Mn and Cu (150?N?+?mix), repeated three times with approximately eight years interval. There was a clear positive effect on volume increment of the 150?N and 150?N?+?mix treatments compared to Control, and the effect was significantly higher for 150?N?+?mix than for 150?N. Fertilization had a stronger effect in the first fertilization period than in the second, while the third period was intermediate. The effect of 150?N?+?mix was strongest at plots?>?300?m a.s.l. However, this correlation may be due to geological conditions rather than elevation. Further studies are needed to find out under which edaphic conditions a nutrient mixture will increase growth substantially in young spruce stands.     

Effects on the flora in Norway spruce forests following clearcutting and shelterwood cutting

Clearcutting and shelterwood cutting of mature Norway spruce peatland forests were compared regarding the effects on the forest flora. Repeated observations of the field and bottom layer vegetation were made before and until 7 or 8 years after harvesting at four sites located along a gradient from southern to northern Sweden. Clearcutting resulted in a greater change of species composition compared with shelterwood cutting. Diversity, measured as Simpson's index, and species number per subplot were lower in the clearcut than in the shelterwood after 7 or 8 years. By applying Ellenberg's indicator values, it was concluded that shelterwood regimes may preserve species preferring shaded and moist conditions, whereas those species decreased after clearcutting. Species preferring high levels of nitrogen increased in the clearcut.According to both multivariate and univariate analyses, the vegetation changed in a similar direction at all sites, although the level of response differed considerably. Despite these similarities, there was a marked site effect, which was not surprising considering the large geographic variation between sites.It is concluded that shelterwood cutting might be a better alternative than clearcutting for forests on fertile peatland sites, with respect to conservation of vascular plants and bryophytes.     

Snow disturbances in secondary Norway spruce forests in Central Europe: Regression modeling and its implications for forest management

Snow is an important ecological factor limiting vegetation distribution, growth, and regeneration, and the importance of snow in the latitudes of Central Europe is expected to increase in the future. We assessed snow damage to secondary spruce stands (not of native provenance therefore not adapted to local conditions) in a mountainous region of 14500 ha in Central Europe (Moravian-Silesian Beskids, Czech Republic). We used neural networks-based regression modeling to study the relationship between stand and environmental parameters and four types of snow damage (top tree, crown, and stem breakage, and uprooting) that occurred during heavy snowfalls in winter 2005/2006 and 2009. Almost 40000 trees were sampled in 345 plots after each of these two events.The results suggest that parameters that can be controlled by forest management (mainly stand density and taper) were not closely associated with spruce forest resistance to snow damage. Investigated snow damage types were primarily related to the developmental stage of the stand, as indicated by stand volume, age, height, and diameter. Damage in 2009, which was caused by shorter-lasting and lower snow loads than the damage in 2005/2006, was also associated with elevation and snow depth. The response of snow damage to all stand development-related variables was clearly unimodal.We infer that forest management can reduce snow damage to secondary spruce forests in Central Europe only to a limited extent, especially under heavy snow loads. This conclusion is supported by the heavy snows that have frequently fallen on forests in Central Europe in the past and the projected increase in winter precipitation in mid- and northern latitudes; thereby increasing snow damage to forest in the future. Therefore, managers of such spruce forests should not specifically consider forest resistance to snow damage but should apply general practices that maintain forest health and productivity.     

Early drought-induced changes to the needle proteome of Norway spruce

To elucidate early drought responses in needles of Norway spruce (Picea abies (Karst.) L.), we subjected 1-year-old seedlings to gradual desiccation for 6 weeks. Four weeks of drought treatment caused a small but significant decrease in photosystem II quantum yield of light-adapted needles (phi(a)) compared with that of well-watered controls. Six weeks of drought treatment reduced phi(a) and the photosystem II quantum yield of dark-adapted needles (phi) by 50 and 8%, respectively, and reduced shoot water potential by 0.7 MPa, but had no measurable effect on needle relative water content. After two weeks of drought treatment, and before there was a discernible effect of drought on phi or a statistically significant effect on shoot water potential, needles were analyzed for changes in protein composition. Five out of several hundred detected proteins in needles of drought-treated plants showed consistent changes compared with control leaves. The proteins were identified by LC-MS/MS as components of the oxygen-evolving complex (oxygen evolving enhancer protein 2), ribulose-1,5-bisphosphate carboxylase/oxygenase large subunit, and one protein of unknown function, whose mRNA was found in a previous screen of wound- and methyl-jasmonate-induced bark proteins.     

Effects of forest restoration treatments on the abundance of bark beetles in Norway spruce forests of southern Finland

Restoration of protected areas in boreal forests frequently includes creating substantial volumes of dead wood. While this benefits a wide range of dead wood dependent invertebrate species, some of these are regarded as forest pests. Therefore, the risk of elevated levels of tree mortality in surrounding commercial forests must be considered. In a large-scale field experiment in southern Finland, we studied the effects of restoration treatments on the abundance of bark beetles within and in the vicinity of restored areas, in particular focusing on Ips typographus and Pityogenes chalcographus. The treatments applied to managed Norway spruce forests were controlled burning and partial harvesting combined with retaining 5, 30 or 60 m³/ha of cut down wood. We found that the abundance of bark beetles increased by both burning and harvesting with down wood retention, being highest where burning and harvesting had been combined. The actual volume of down wood retention had no significant effect. The effect of burning on the number of bark beetles along host tree boles was negative which suggests that burnt spruces provided a less suitable resource for bark beetles than unburnt dead spruces. The abundance of bark beetles along host trees also decreased with increasing volume of down wood retention. The abundance of P. chalographus was slightly elevated up to 50 m outside restored areas but the abundance was very low compared to that within the areas. The abundance of I. typographus was extremely low outside restored areas. We conclude that restoration treatments increase the abundance of bark beetles via increased availability of resources, but that the effect of burning is likely to be counteracted by decreased resource quality. Thus, burning might be the “safest” way to produce large quantities of dead wood. Furthermore, the fact that only few beetles were collected in adjacent areas suggests that restored areas pose little threat of serving as refugia in which bark beetle populations increase in sufficient numbers to attack live trees in adjacent forests. However, restoration actions repeated at consecutive years within a small area might enable the populations to grow to outbreak levels.     

Effects of elevated CO(2) concentration and nutrition on net photosynthesis,stomatal conductance and needle respiration of field-grown Norway spruce trees

To study the effects of elevated CO(2) on gas exchange, nonstructural carbohydrate and nutrient concentrations in current-year foliage of 30-year-old Norway spruce (Picea abies (L.) Karst.) trees, branches were enclosed in ventilated, transparent plastic bags and flushed with ambient air (mean 370 &mgr;mol CO(2) mol(-1); control) or ambient air + 340 &mgr;mol CO(2) mol(-1) (elevated CO(2)) during two growing seasons. One branch bag was installed on each of 24 selected trees from control and fertilized plots. To reduce the effect of variation among trees, results from each treated branch were compared with those from a control branch on the same whorl of the same tree. Elevated CO(2) increased rates of light-saturated photosynthesis on average by 55% when measured at the treatment CO(2) concentration. The increase was larger in shoots with high needle nitrogen concentrations than in shoots with low needle nitrogen concentrations. However, shoots grown in elevated CO(2) showed a decrease in photosynthetic capacity compared with shoots grown in ambient CO(2). When measured at the internal CO(2) concentration of 200 &mgr;mol CO(2) mol(-1), photosynthetic rates of branches in the elevated CO(2) treatments were reduced by 8 to 32%. The elevated CO(2) treatment caused a 9 to 20% reduction in carboxylation efficiency and an 18% increase in respiration rates. In response to elevated CO(2), starch, fructose and glucose concentrations in the needles increased on average 33%, whereas concentrations of potassium, nitrogen, phosphorus, magnesium and boron decreased. Needle nitrogen concentrations explained 50-60% of the variation in photosynthesis and CO(2) acclimation was greater at low nitrogen concentrations than at high nitrogen concentrations. We conclude that the enhanced photosynthetic rates found in shoots exposed to elevated CO(2) increased carbohydrate concentrations, which may have a negative feedback on the photosynthetic apparatus and stimulate cyanide-resistant respiration. We also infer that the decrease in nutrient concentrations of needles exposed to elevated CO(2) was the result of retranslocation of nutrients to other parts of the branch or tree.     

The chemical composition of needle and leaf litter from Scots pine, Norway spruce and white birch in Scandinavian forests

Needle litter from 14 stands of Scots pine (Pinus silvestris,L.), 13 stands of Norway spruce (Picea abies (L.) Karst.) andleaf litter from three stands of white birch (Betula pubescensEhrh.) were analysed for chemical composition. The concentrationsof the elements N, P, K, Ca, Mg and Mn as well as solid organiccomponents (lignin, cellulose and hemicelluloses) and solubleswere determined. When the average chemical compositions werecompared the Scots pine needle litter was clearly the most nutrient-poorlitter type. Of the solid organic-chemical components the ligninfraction dominated in the spruce and birch litter whereas thecellulose dominated in the pine needle litter. When Norway spruce and Scots pine were growing in adjacent standson soils with the same bedrock origin the spruce litter hadsignificantly higher concentrations of nutrients (N, P, K, Ca,Mg, Mn) than the pine needle litter. At sites where Norway spruceand white birch were growing in adjacent stands, the birch leaflitter had generally higher concentrations of nutrients. However, significant or nearly significant differences were onlyobtained for Mg (P = 0.002), K (P = 0.056) and N (P = 0.087),probably due to the few replicates of stands compared. Concerningorganic chemical components, the spruce needle litter had significantlyhigher concentrations of lignin and mannan than all the otherlitters and lower levels of ethanol-soluble substances, celluloseand galactan than the pine needle litter. Further, it had lowerconcentrations of water solubles, rhamnan and xylan than thebirch litter. No relationships were established between the nutrient statusof the conifer litters and the site index H100 (the dominantheight of the trees at a reference age of 100 years) of thestands. Concentrations of solid carbohydrates in the litterswere, however, positively correlated with site index (P <0.001). Further, the concentration of nitrogen in the pine needlelitter was negatively correlated with the latitude of the sites(P < 0.01). The influence of litter chemistry on the decompositionof litter and nutrient cycling of forests is discussed.     

Effects of planting time on pine weevil (Hylobius abietis) damage to Norway spruce seedlings

Feeding by pine weevil (Hylobius abietis L.) causes severe damage to newly planted conifer seedlings in most parts of Scandinavia. We investigated the effect of planting time and insecticide treatment on pine weevil damage and seedling growth. The main objective was to study if planting in early autumn on fresh clear-cuts would promote seedling establishment and reduce the amount of damage caused by pine weevil the following season. The experiment was conducted in southern Sweden and in south-eastern Norway with an identical experimental design at three sites in each country. On each site, Norway spruce seedlings with or without insecticide treatment were planted at four different planting times: August, September, November and May the following year. In Sweden, the proportion of untreated seedlings that were killed by pine weevils was reduced when seedlings were planted at the earliest time (August/September) compared to late planting in November, or May the following year. This pattern was not found in Norway. The average length of leading shoot, diameter growth and biomass were clearly benefited by planting in August in both countries. Insecticide treatment decreased the number of seedlings killed or severely damaged in both Norway and Sweden.     

Effects of photoperiod and temperature on the timing of bud burst in Norway spruce (Picea abies)

We examined the effects of several photoperiod and temperature regimes imposed during the winter-spring period on the timing of bud burst in rooted cuttings of Norway spruce (Picea abies (L.) Karst.) grown in a greenhouse in Finland. The treatments were initiated in November and December after the cuttings had been exposed to natural chilling and freezing events. Irrespective of the treatments applied, time to bud burst decreased with increased duration of previous exposure to natural chilling and freezing events. Fluctuating day/night temperatures and continuous lengthening of the photoperiod hastened bud burst. Shortening the photoperiod delayed bud burst, suggesting that little or no ontogenetic development toward bud burst takes place during mild periods before the winter solstice. In the case of climatic warming, this phenomenon may prevent the premature onset of growth that has been predicted by computer simulations with models that only consider temperature regulation of bud burst.     

Polyamines in embryogenic cultures of Norway spruce (Picea abies) and red spruce (Picea rubens)

Embryogenic cultures of red spruce (Picea rubens Sarg.) and Norway spruce (Picea abies (L.) Karst.) were initiated from dissected mature zygotic embryos. The tissues were grown on either proliferation medium or maturation medium. On proliferation medium, the embryogenic tissue continued to produce early stage somatic embryos (organized meristems attached to elongated, suspensor-like cells), whereas on maturation medium fully mature embryos developed from the embryonic tissue. Analysis of polyamines in tissues grown on these two media showed that: (1) both putrescine and spermidine concentrations were always higher in cultures grown on proliferation medium than in cultures grown on maturation medium; (2) in both species, spermidine concentrations declined with time in the tissues grown on maturation medium; and (3) spermine was present in only minute quantities and showed only a small change with time. The presence of difluoromethylornithine in the culture medium had little effect on polyamine concentration, whereas the presence of difluoromethylarginine caused a decrease in putrescine concentrations in both red spruce and Norway spruce tissues grown on proliferation medium or maturation medium.     

Conversion of Norway spruce forests in the face of climate change: a case study in Central Europe

Steadily increasing damage to Norway spruce forests in Europe has caused researchers and managers to consider whether these forests can be converted to more stable ecosystems. In a central European mountain region, we investigated whether management systems (MSs) specified by regional stakeholders provide sound alternatives to the currently applied management. We used the forest model Sibyla to explore whether the tested MSs differ in their sensitivity to climate change in terms of altered biomass production, stand structure, forest damage, and financial outcome. The tested MSs were no-management (NM), currently applied management (BAU), and management based on the preferences of forest managers (FM) or on the preferences of other stakeholders (OSH). With NM, spruce remained dominant during the simulation period 2010–2100, and the rate of damage significantly increased. Spruce also remained dominant with FM, while the abundance of non-spruce species significantly increased with BAU and OSH. The rate of salvage logging converged at 50% of the total harvest for all MSs up to 2050. Climate change reduced biomass production (?15%) with all MSs but had a negligible effect on biodiversity indicators. The average initial value of the simulated stands was 20,000 € ha^?1 and the nominal value in 2100 was between 1900 and 10,900 € ha^?1. The Net Present Value calculated with the 2% interest rate was negative during the whole simulation period (?5600 to ?18,500 € ha^?1 in 2100). Effect of climate change on all financial indicators was negative. Our findings indicate that secondary spruce forests are highly vulnerable and that the systems proposed by both forest managers and other regional stakeholders failed to significantly reduce forest damage and stabilize forest production.     

Persisting bark beetle outbreak indicates the unsustainability of secondary Norway spruce forests: case study from Central Europe

? Context

Secondary Norway spruce forests in the Western Beskids are among the most damaged forests in Europe. Although spruce bark beetle (Ips typographus) has been recently causing large-scale damage to these forests, our understanding of I. typographus dynamics in this environment is inadequate for evaluating forest sustainability.

? Aim

This study aims to evaluate the patterns of damage caused by I. typographus to spruce forests with compromised ecological stability.

? Methods

Forest infestation by I. typographus was inferred from sanitary felling data collected from 1998 to 2004. Stand and site data were obtained from forest management plans. Spatial-dependence analysis, ordinary kriging and neural network-based regression modelling were used to investigate the patterns of infestation and the casual relationships in the studied ecosystem.

? Results

I. typographus long-distance dispersal substantially decreased with outbreak culmination. The spread of infestation was only weakly related to stand and site parameters. Infestations spread isotropically at the stand and patch level but directionally at the regional scale.

? Conclusions

The large-scale spread of infestation can be explained by the uniform age and species composition of the investigated forests and by the ability of populations to overwhelm suboptimal trees. The observations presented here suggest that secondary spruce forests in Europe may be unsustainable due to unprecedented bark beetle outbreaks, which can be further amplified by changing climate.     

Effects of experimental drought on the fine root system of mature Norway spruce

Norway spruce (Picea abies (L.) Karst.) is an economically important, but relatively drought-sensitive tree species that might suffer from increasing drought intensities and frequencies, which are predicted to occur in parts of central Europe under future climatic change. In a throughfall exclusion experiment using sub-canopy roofs, we tested the hypothesis that enhanced drought leads to an increase in fine root mortality, and also to a higher, subsequent fine root growth. Fine root production and mortality were assessed using two independent approaches, sequential soil coring (organic layer) and direct observations in minirhizotrons (organic layer plus upper mineral soil). Six weeks of throughfall exclusion resulted in mild drought stress, i.e. a reduction in average soil moisture from 20 to 12 vol.% during the treatment. Based on the sequential coring data, experimental drought did not result in significant changes in fine root biomass during the 6-week treatment period, but caused an increase in fine root mortality by 61% in the 6 weeks following the drought treatment. Remarkably, fine root production showed a synchronous increase in this period, which more than compensated for the loss due to increased mortality. The minirhizotron data confirmed that the drought treatment increased fine root loss in the organic layer. Based on this method, however, root loss occurred during the drought period and was not compensated by increased root production. The mild drought stress was mainly restricted to the organic layer and did not significantly influence fine root dynamics in the mineral soil. We calculated that the drought event resulted in an extra input of about 28 g C m⁻² and 1.1 g N m⁻² to the soil due to increased fine root mortality. We conclude that even periods of mild drought significantly increase fine root mortality and the associated input of root-derived C to the soil organic matter pool in temperate Norway spruce forests.     

Effects of cone-induction treatments on black spruce (Picea mariana) current-year needle development and gas exchange properties

Both drought and root pruning (RP) increased the number of cones induced when black spruce (Picea mariana (Mill.) B.S.P.) grafts were injected with gibberellins A(4/7) (GA), but their effects on predawn shoot water potential and current-year needle development differed. Drought decreased predawn shoot water potential (Psi(pd)), but only during the period when irrigation was withheld, and it had no effect on the growth or gas exchange properties of current-year needles. Conversely, root pruning had little effect on Psi(pd), but it resulted in trees with smaller current-year needles that had lower nitrogen and chlorophyll concentrations and reduced rates of gas exchange up to the later stages of shoot elongation compared with needles of control trees. These findings are discussed in relation to potential effects on the development of induced cones in the following growth cycle.