Different responses of northern and southern ecotypes of Betula pendula to exogenous ABA application

We investigated responses of northern and southern ecotypes of silver birch (Betula pendula Roth) to exogenous abscisic acid (ABA) under controlled environmental conditions to determine the role of ABA in cold acclimation and dormancy development. Abscisic acid was sprayed on the leaves and changes in freezing tolerance, determined by the electrolyte leakage test, and bud dormancy were monitored. Applied ABA induced cold acclimation but had no effect on growth cessation in seedlings grown in long day conditions (LD, 24-h photoperiod at 18 degrees C). It enhanced freezing tolerance and accelerated growth cessation in seedlings grown in short day conditions (SD, 12-h photoperiod at 18 degrees C), and slightly enhanced freezing tolerance in seedlings grown at low temperature (LT, 24-h photoperiod at 4 degrees C) in both ecotypes. There were distinct ecotypic differences in ABA-induced cold acclimation and dormancy development. The northern ecotype was more responsive to applied ABA than the southern ecotype, resulting in more rapid development of freezing tolerance in all treatments, and earlier dormancy development in SD. When plants were grown in a photoperiod just above the critical photoperiod for the ecotype (defined as the longest photoperiod that induces growth cessation), applied ABA caused growth cessation and dormancy development. Compared with ABA-treated seedlings grown in SD, dormancy development was delayed in ABA-treated seedlings exposed to a near-critical photoperiod, but even in this treatment dormancy developed faster in the northern ecotype than in the southern ecotype.     

High autumn temperature delays spring bud burst in boreal trees, counterbalancing the effect of climatic warming

The effect of temperature during short-day (SD) dormancy induction was examined in three boreal tree species in a controlled environment. Saplings of Betula pendula Roth, B. pubescens Ehrh. and Alnus glutinosa (L.) Moench. were exposed to 5 weeks of 10-h SD induction at 9, 15 and 21 degrees C followed by chilling at 5 degrees C for 40, 70, 100 and 130 days and subsequent forcing at 15 degrees C in a 24-h photoperiod for 60 days. In all species and with all chilling periods, high temperature during SD dormancy induction significantly delayed bud burst during subsequent flushing at 15 degrees C. In A. glutinosa, high temperature during SD dormancy induction also significantly increased the chilling requirement for dormancy release. Field experiments at 60 degrees N with a range of latitudinal birch populations revealed a highly significant correlation between autumn temperature and days to bud burst in the subsequent spring. September temperature alone explained 20% of the variation between years in time of bud burst. In birch populations from 69 and 71 degrees N, which ceased growing and shed their leaves in August when the mean temperature was 15 degrees C, bud burst occurred later than expected compared with lower latitude populations (56 degrees N) in which dormancy induction took place more than 2 months later at a mean temperature of about 6 degrees C. It is concluded that this autumn temperature response may be important for counterbalancing the potentially adverse effects of higher winter temperatures on dormancy stability of boreal trees during climate warming.     

Climatic control of bud burst in young seedlings of nine provenances of Norway spruce

Detailed knowledge of temperature effects on the timing of dormancy development and bud burst will help evaluate the impacts of climate change on forest trees. We tested the effects of temperature applied during short-day treatment, duration of short-day treatment, duration of chilling and light regime applied during forcing on the timing of bud burst in 1- and 2-year-old seedlings of nine provenances of Norway spruce (Picea abies (L.) Karst.). High temperature during dormancy induction, little or no chilling and low temperature during forcing all delayed dormancy release but did not prevent bud burst or growth onset provided the seedlings were forced under long-day conditions. Without chilling, bud burst occurred in about 20% of seedlings kept in short days at 12 degrees C, indicating that young Norway spruce seedlings do not exhibit true bud dormancy. Chilling hastened bud burst and removed the long photoperiod requirement, but the effect of high temperature applied during dormancy induction was observed even after prolonged chilling. Extension of the short-day treatment from 4 to 8 or 12 weeks hastened bud burst. The effect of treatments applied during dormancy development was larger than that of provenance; in some cases no provenance effect was detected, but in 1-year-old seedlings, time to bud burst decreased linearly with increasing latitude of origin. Differences among provenances were complicated by different responses of some origins to light conditions under long-day forcing. In conclusion, timing of bud burst in Norway spruce seedlings is significantly affected by temperature during bud set, and these effects are modified by chilling and environmental conditions during forcing.     

Maturation of maritime pine (Pinus pinaster Ait.) seedlings after exposure to a period of continuous light

Nine half-sib families of maritime pine (Pinus pinaster Ait.) of known adult performance were grown in continuous light at either 25 degrees C or 25/20 degrees C for 18 weeks. They were then exposed to a dormancy induction period followed by a dormancy release period and then grown for a further 9 weeks in a 16-h photoperiod at a day/night temperature of 25/20 degrees C. Seedlings exhibited great diversity in morphology at the end of the first growth period. The number of morphogenetic cycles varied between one and three and the form of the apical meristem ranged from a typical rosette to an adult-like bud. The type of seedling obtained at the end of the first growth period strongly influenced later growth, independently of the temperature regime. Maturity was proportional to the number of morphogenetic cycles achieved during the first growth period and was characterized by short growth duration, small primary needles and a high degree of fixed growth. The state of the apical meristem that underwent the dormancy period had less influence on the rate of maturation than the number of morphogenetic cycles. The time course of maturation was endogenously controlled and varied among traits. Conspicuous morphological differences were not associated with changes in the relationship between growth components at the phenotypic level. However, there seemed to be a shift in the genetic correlations between growth components after first budset.     

The effect of different atmospheric ozone partial pressures on photosynthesis and growth of nine fruit and nut tree species

Nursery stock of peach (Prunus persica L. Batsch, cv. O'Henry), nectarine (P. persica L. Batsch, cv. Fantasia), plum (P. salicina Lindel., cv. Casselman), apricot (P. armeniaca L., cv. Tilton), almond (P. dulcis Mill., cv. Nonpareil), prune (P. domestica L., cv. Improved French), cherry (P. avium L., cv. Bing), oriental pear (Pyrus pyrifolia Rehd., cv. 20th Century), and apple (Malus pumula Mill., cv. Granny Smith) were planted in open-top chambers on April 1, 1988 at the University of California's Kearney Agricultural Center located in the San Joaquin Valley (30 degrees 40' N 119 degrees 40' W). Trees were exposed to three atmospheric ozone partial pressures (charcoal-filtered air (C), ambient air (A), or ambient air + ozone (T)) from August 1 to November 17, 1988. The mean 12-h (0800 to 2000 h) ozone partial pressures measured in open-top chambers during the experimental period were 0.030, 0.051, and 0.117 microPa Pa(-1) in the C, A and T treatments, respectively. Leaf net CO(2) assimilation rate decreased linearly with increasing 12-h mean ozone partial pressure for the almond, plum, apricot, prune, pear, and apple cultivars. Stomatal conductances of apricot, apple, almond, and plum decreased linearly with increasing ozone partial pressure. Cross-sectional area relative growth rates of almond, plum, apricot, and pear decreased linearly with increasing ozone partial pressure. Net CO(2) assimilation rate, stomatal conductance, and trunk growth of cherry, peach and nectarine were unaffected by the ozone treatments. Reduced leaf gas exchange probably contributed to ozone-induced growth reduction of the susceptible species and cultivars. Several of the commercial fruit tree species and cultivars studied were relatively tolerant to the ozone treatments.     

Effect of Temperature on the Induction of Bud Dormancy in Ecotypes of Betula pubescens and Betula pendula

The purpose of this study was to determine the influence of temperature applied during short day-induced budset on induction of dormancy in six ecotypes of Betula pubescens Ehrh. and two ecotypes of Betula pendula Roth. Seedlings were grown in a phytotron at constant temperatures of 9–21°C under a 12 h photoperiod (SD) during dormancy induction. Induction of dormancy was monitored by following bud flushing and shoot growth after transfer to long photoperiod conditions (24 h) at 18°C. Chilling requirement was studied in seedlings exposed to 10 weeks of SD. In both species induction of bud dormancy developed most rapidly at 15–18°C, and both 9–12°C and 21°C delayed the induction of dormancy. Raising the temperature (from 9 to 21°C) applied during induction of dormancy significantly increased the chilling requirement. These responses were noted for all ecotypes tested, but in general the northern ecotypes entered dormancy more quickly than the southern ones. No such trend was recorded for chilling requirement, although a B. pubescens ecotype from Iceland and another from the coast of northern Norway appeared to require a longer chilling treatment than the other ecotypes. In conclusion, induction and depth of bud dormancy in birch are significantly affected by temperature conditions and these effects may explain some of the annual variation in dormancy and chilling requirement observed in nature.     

Flooding, root temperature, physiology and growth of two Annona species

The effects of root zone temperature (RZT) and flooding on physiology and growth of Annona glabra L. (pond apple) and A. muricata L. (soursop) were investigated. Trees planted in containers were exposed to RZTs of 5, 10, 20, 25 or 35 degrees C in controlled root temperature chambers. Trees at each RZT were either non-flooded (control) or continuously flooded. There were four replications over time for each treatment combination. Pond apple was more flood-tolerant than soursop. A combination of flooding and RZTs of 5 and 10 degrees C resulted in tree mortality of both species by Week 4. Only trees that appeared to develop morphological adaptations survived continuous flooding. In both species, net CO2 assimilation (A) decreased to nearly zero within 1 week following exposure to RZTs of 5 or 10 degrees C and became consistently negative over the remaining experimental period. Flooding reduced leaf chlorophyll index (measured with a SPAD meter), A and plant growth, and increased root electrolyte leakage from soursop. Optimum growth occurred at RZTs of 25 to 35 degrees C for non-flooded pond apple trees and at 20 to 25 degrees C for flooded trees. Soursop exhibited maximum growth at RZTs of 35 degrees C under non-flooded conditions and at 25 degrees C under flooded conditions.     

Differential responses of silver birch (Betula pendula) ecotypes to short-day photoperiod and low temperature

We investigated interrelations of dormancy and freezing tolerance and the role of endogenous abscisic acid (ABA) in the development of silver birch (Betula pendula Roth) ecotypes in controlled environments. Short-day treatment induced growth cessation, bud set and dormancy development, as well as initiation of cold acclimation and an increase in freezing tolerance. Subsequent low temperature and short days (12-h photoperiod) resulted in a significant increase in freezing tolerance, whereas bud dormancy was gradually released. The concentration of ABA increased in response to short days and then remained high, but ABA concentrations fluctuated irregularly when the dormant plants were subsequently exposed to low temperature during short days. Although there was a parallel development of freezing tolerance and bud dormancy in response to short days, subsequent exposure to low temperature had opposite effects on these processes, enhancing freezing tolerance and releasing dormancy. Compared with the southern ecotype, the northern ecotype was more responsive to short days and low temperature, exhibiting earlier initiation of cold acclimation, growth cessation and an increase in ABA concentrations in short days, and higher freezing tolerance, faster dormancy release and greater alteration in ABA concentrations when subsequently exposed to low temperature during short days. The rates and extent of the increases in ABA concentration may be related to increases in freezing tolerance and dormancy development during short days, whereas the extent of the fluctuations in ABA concentration may play an important role in enhancing freezing tolerance and releasing dormancy during a subsequent exposure to low temperature during short days.     

Effects of ABA application on cessation of shoot elongation in long-day grown Norway spruce seedlings

Abscisic acid (ABA) was applied in lanolin to apical buds of Norway spruce (Picea abies (L.) Karst.) seedlings actively growing in a 24 h photoperiod. At a rate of 100 microg per plant, ABA suspended shoot elongation for about three weeks in the majority of plants but failed to induce normal winter buds. The role of ABA in the induction of dormancy is thus uncertain in conifers as well as in deciduous woody plants.     

Root-zone temperatures affect phenology of bud break, flower cluster development, shoot extension growth and gas exchange of 'Braeburn' (Malus domestica) apple trees

We investigated the effects of root-zone temperature on bud break, flowering, shoot growth and gas exchange of potted mature apple (Malus domestica (Borkh.)) trees with undisturbed roots. Soil respiration was also determined. Potted 'Braeburn' apple trees on M.9 rootstock were grown for 70 days in a constant day/night temperature regime (25/18 degrees C) and one of three constant root-zone temperatures (7, 15 and 25 degrees C). Both the proportion and timing of bud break were significantly enhanced as root-zone temperature increased. Rate of floral cluster opening was also markedly increased with increasing root-zone temperature. Shoot length increased but shoot girth growth declined as root-zone temperatures increased. Soil respiration and leaf photosynthesis generally increased as root-zone temperatures increased. Results indicate that apple trees growing in regions where root zone temperatures are < or = 15 degrees C have delayed bud break and up to 20% fewer clusters than apple trees exposed to root zone temperatures of > or = 15 degrees C. The effect of root-zone temperature on shoot performance may be mediated through the mobilization of root reserves, although the role of phytohormones cannot be discounted. Variation in leaf photosynthesis across the temperature treatments was inadequately explained by stomatal conductance. Given that root growth increases with increasing temperature, changes in sink activity induced by the root-zone temperature treatments provide a possible explanation for the non-stomatal effect on photosynthesis. Irrespective of underlying mechanisms, root-zone temperatures influence bud break and flowering in apple trees.     

Autumn temperature affects the induction of dormancy in first‐year seedlings of acer platanoides L.

First‐year seedlings of five latitudinal populations of Acer platanoides were subjected to decreasing photoperiod treatment under three different temperature regimes. The depth of the induced dormancy was quantified as the number of days to bud burst (DBB) under defined conditions favourable to growth. The results suggested a close relationship between autumn temperature and the strength of the induced dormancy, with high temperatures combined with short days leading to a deeper stage of dormancy. Northern and continental populations generally had bud burst earlier than southern. The results are discussed in relation to hypotheses for dormancy induction and release.     

Relationships among cold hardiness, root growth potential and bud dormancy in three conifers

Greenhouse-cultured, container-grown ponderosa pine (Pinus ponderosa var. scopulorum Engelm.), interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) and Engelmann spruce (Picea engelmannii (Parry) Engelm.) were cold acclimated and deacclimated in growth chambers over 19 weeks. Stem cold hardiness, total new root length at 14 days and days to bud break were measured weekly. Relationships among cold hardiness, root growth potential (RGP) and bud dormancy suggest that cold hardiness, which can be measured quickly, could provide a useful basis for estimating the two other parameters. During cold acclimation, there was a lag period in which stem cold hardiness remained at -15 degrees C and RGP was at a minimum, in all three species. Douglas-fir and Engelmann spruce buds remained fully dormant during this lag period. Ponderosa pine buds had no chilling requirement for the loss of dormancy, and reached quiescence during the lag period. Immediately following the lag period, as stem cold hardiness progressed to -22 degrees C, RGP increased to a high plateau in all three species, and Douglas-fir and Engelmann spruce buds approached quiescence. Cold deacclimation and bud development began immediately on exposure to warm, long days, but RGP remained high until stem cold hardiness returned to approximately -15 degrees C. At bud break, cold hardiness and RGP were at the minimum.     

A molecular marker associated with low-temperature induction of dormancy in red osier dogwood (Cornus sericea)

Dormancy induction in temperate deciduous plants is thought to be regulated by short photoperiods, but low temperature has been shown to eliminate the short photoperiod requirement in northern ecotypes. An F2 population (191 plants) red osier dogwood (Cornus sericea L.) derived from a polycross of an F1 population produced from reciprocal crosses of the parental clonal ecotypes, Northwest Territories (NWT, 62 degrees N) and Utah (42 degrees N), was examined to identify molecular markers of temperature-induced endodormancy. Dormancy induction curves were generated for each individual in the F2 population and a standard point prior to vegetative maturity (i-VM) was inferred from the change in slope of the dormancy acquisition curve. Under Saskatoon, Saskatchewan field conditions (52 degrees N), the NWT ecotype entered i-VM on average 5-6 weeks before the Utah ecotype. Two sub-populations of the F2 population were distinguishable based on VM acquisition on exposure to low temperature but not to short photoperiods. A sequence characterized amplified region (SCAR) marker was developed that correctly (> 92%) identified individual plants within the F2 subpopulation that were responsive to low-temperature induction of VM. Timing of bud break was strongly associated with the timing of VM in the geographical ecotypes but not in the F2 population, indicating that these are separate traits under genetic control.     

Effects of "near-lethal" stress on bud dormancy and stem cold hardiness in red-osier dogwood

We studied the effects of "near-lethal" (NL, 47 degrees C for 1 h) heat stress, applied to intact shoots of red-osier dogwood (Cornus sericea L.) during early (October), deep (November) or late (December) dormancy, on bud dormancy release and development of stem tissue cold hardiness under natural conditions and at a constant temperature of 0 or 23 degrees C in the dark. The NL heat-stress treatment overcame bud dormancy when applied during the early and late stages of dormancy. During October and December, all plants in the 23 degrees C + dark post-stress environment broke bud within 35 and 12 days, respectively, whereas the corresponding values for days to bud break in the control plants were more than 150 and 110 days, respectively. Application of NL heat stress during deep dormancy caused only slightly earlier bud break compared to the control plants. In the 0 degrees C + dark post-stress environment, all NL heat-treated plants died within 9 weeks. Under natural post-stress conditions, bud break in plants receiving NL heat stress during early and deep dormancy occurred at the same time as in control plants, whereas bud break of plants receiving NL heat stress during late dormancy occurred 55 days earlier than in control plants. Under both natural and 23 degrees C + dark post-stress conditions, cold hardiness of plants receiving NL heat stress during early dormancy was similar to that of controls. Application of NL heat stress during deep dormancy hastened the rate of deacclimation under the 23 degrees C + dark post-stress conditions but had no effect on deacclimation under natural post-stress conditions. Application of NL heat stress during late dormancy enhanced deacclimation of plants in both the 23 degrees C + dark and natural post-stress environments.     

Dormancy release and chilling requirement of buds of latitudinal ecotypes of Betula pendula and B. pubescens

Bud burst and dormancy release of latitudinal ecotypes of Betula pendula Roth and B. pubescens Ehrh. from Denmark ( approximately 56 degrees N), mid-Norway ( approximately 64 degrees N) and northern Norway ( approximately 69 degrees N) were studied in controlled environments. Dormant seedlings were chilled at 0, 5 or 10 degrees C from October 4 onward and then, at monthly intervals from mid-November to February, batches of seedlings were held at 15 degrees C in an 8-h (SD) or 24-h (LD) photoperiod to permit flushing. A decline in days to bud burst occurred with increasing chilling time in all ecotypes. In November, after 44 chilling days, time to bud burst was least in plants chilled at 0 and 5 degrees C. The difference diminished with increasing chilling time, and in February, after 136 chilling days, bud burst was earliest in plants chilled at 10 degrees C. Long photoperiods during flushing significantly reduced thermal time after short chilling periods (44 and 74 days), but had no effect when the chilling requirement was fully met after 105 or more chilling days. No significant difference in these responses was found between the two species. In both species, chilling requirement decreased significantly with increasing latitude of origin. Bud burst was normal in seedlings overwintered at 12 degrees C, but was erratic and delayed in seedlings overwintered at 15 and especially at 21 degrees C, indicating that the critical overwintering temperature is between 12 and 15 degrees C. We conclude that there is little risk of a chilling deficit in birch under Scandinavian winter conditions even with a climatic warming of 7-8 degrees C. The likely effects of a climatic warming include earlier bud burst, a longer growing season and increased risk of spring frost injury, especially in high latitude ecotypes.     

Influence of photoperiod and temperature on frost hardiness and free amino acid concentrations in black spruce seedlings

We investigated the effects of a 4-week exposure to an 8-h or 18-h photoperiod at 5 or 25 degrees C on the development of hardiness to -20 degrees C and the accumulation of proline (Pro), arginine (Arg) and tryptophan (Trp) in shoots of black spruce (Picea mariana (Mill.) B.S.P.) seedlings. The greatest degree of hardening to -20 degrees C occurred in seedlings exposed to an 8-h photoperiod at 25 degrees C, and some hardening occurred in seedlings exposed to 5 degrees C in either an 8-h or 18-h photoperiod. Proline accumulated in shoots in response to 5 degrees C and either an 8-h or 18-h photoperiod, whereas Trp accumulated in response to an 8-h photoperiod at either temperature, and Arg only accumulated in shoots in the 5 degrees C + 8-h photoperiod treatment. Only the accumulation of Trp was significantly related to the degree of hardiness to -20 degrees C.     

Seed dormancy and germination of Ficus lundellii and tropical forest restoration

We investigated seed dormancy and germination in Ficus lundellii Standl. (Moraceae), a native species of Mexico's Los Tuxtlas tropical rain forest. In an 8-h photoperiod at an alternating diurnal (16/8 h) temperature of 20/30 degrees C, germination was essentially complete (96%) within 28 days, whereas in darkness, all seeds remained dormant. Neither potassium nitrate (0.05-0.2%) applied continuously nor gibberellic acid applied either continuously (10-200 ppm) or as a 24 hour pretreatment (2000 ppm) induced germination in the dark. Germination in the light was not reduced by a 24-h hydrochloric acid (0.1-1%) pretreatment, but it was reduced both by a 24-h pretreatment with either H(2)O(2) (0.1-5 M) or 5% HCl, or by more than 5 days of storage at 40 degrees C (4.5% seed water content). In a study with a 2-dimensional temperature gradient plate, seeds germinated fully and rapidly in the light at a constant temperature of 30 degrees C, and fully but less rapidly in the light at alternating temperatures with low amplitudes (< 12 degrees C) about the optimal constant temperature. The base, optimal and ceiling temperatures for rate of germination were estimated as 13.8, 30.1 and 41.1 degrees C, respectively. In all temperature regimes, light was essential for the germination of F. lundellii seeds.     

The biological efficacy of pear ester on the activity of Granulosis virus for codling moth

Ethyl (E,Z)-2,4-decadienoate (pear ester) is an adult and larval kairomonal attractant for Cydia pomonella (L.) (Lepidoptera: Tortricidae). The possibility of using a microencapsulated formulation of pear ester (DA-MEC, a.i. 5%, Trécé Inc.) to interfere with the host location behaviour was evaluated. Laboratory leaf disc bioassays and field efficacy trials were carried out on apple to determine the potential of improving the insecticidal performance of the granulovirus of C. pomonella (CpGV) by co-mixing with pear ester in a sprayable formulation. In laboratory bioassays, adding DA-MEC at low doses of CpGV insecticide increased the larval mortality as a function of time. The field tests performed revealed a significant effect of the blank DA-MEC formulation in reducing fruit injury compared to an unsprayed control.     

Growth and survival of seedlings of various picea species under northern climatic conditions

The main purpose of these studies was to investigate the adaptation of young seedlings of various seed lots of Picea abies (L.) Karst., P. glauca (Moench) Voss, P. x lutzii Little and P. sitchensis (Bong.) Carr. to northern climatic conditions. Effects of temperature and photoperiod on elongation growth and growth cessation were studied under controlled conditions in a phytotron. In addition, growth and survival of the seedlings outdoors at 69°39’ N lat. were followed for two years. Seed lots of /’. abies originated from northern Norway (66° N lat.), those of the other species were from Alaska (between 57°.and 66° N lat.). The critical photoperiod for budset was 19–20 h for seed lots of P. abies. In general, the critical photoperiod for budset increased with increasing latitude of the seed source, but the results indicated a significantly shorter cirtical photoperiod for seed lots from about 60° N lat., <120 m a.s.l. of P. sitchensis (between 12 and 16 h) than for comparable seed lots of P x lutzii (17–18 h) or P. glauca (18 h). The time course of budset under natural light conditions, both in the phytotron and outdoors, generally followed the pattern predicted from the critical photoperiod. However, in P. glauca the budset occurred earlier than in P. abies although the latter had a longer critical photoperiod. Due to the short critical photoperiod and consequently delayed growth cessation and hardening, all seed lots of P. sitchensis (from 58° to 60° N lat.) were severely damaged during winter. Some damage was also observed in P. x lutzii and in P. abies. The optimum temperature for elongation growth was higher for P. sitchensis than for the other species. In the phytotron experiments, seedlings of P. sitchensis grew best at temperatures between 12 and 21°C, but at 9°C the best growth was obtained in some seed lots of P. abies. After two growth seasons outdoors, all seed lots of P. abies were taller than any seed lot of the other species. Also P. glauca seed lots and one seed lot of P. x lutzii showed good growth, and their growth rhythm seemed to be well adapted to the northern conditions.     

Hydraulic architecture and photosynthetic capacity as constraints on release from suppression in Douglas-fir and western hemlock

We compared hydraulic architecture, photosynthesis and growth in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), a shade-intolerant species, and western hemlock (Tsuga heterophylla (Raf.) Sarg.), a shade-tolerant species, to study the temporal pattern of release from suppressive shade. In particular, we sought to determine whether hydraulic architecture or photosynthetic capacity is most important in constraining release. The study was conducted at two sites with mixed stands of 10- to 20-year-old Douglas-fir and western hemlock. At one site, the stand had been thinned allowing release of the understory trees, whereas at the other site, the stand remained unthinned. Douglas-fir had lower height growth (from 1998-2003) and lower relative height growth (height growth from 1998 to 2003/height in 1998) than western hemlock. However, relative height growth of released versus suppressed trees was higher in Douglas-fir (130%) than in western hemlock (65%), indicating that, although absolute height growth was less, Douglas-fir did release from suppression. Release seemed to be constrained initially by a limited photosynthetic capacity in both species. Five years after release, Douglas-fir trees had 14 times the leaf area and 1.5 times the leaf nitrogen concentration (N (area)) of suppressed trees. Needles of released western hemlock trees had about twice the maximum assimilation rate (A (max)) at ambient [CO(2)] as needles of suppressed trees and exhibited no photoinhibition at the highest irradiances. After release, trees increased in leaf area, leaf N concentration and overall photosynthetic capacity. Subsequently, hydraulic architecture appeared to constrain release in Douglas-fir and, to a lesser extent, in western hemlock. Released trees had significantly less negative foliar delta(13)C values than suppressed trees and showed a positive relationship between leaf area:sapwood area ratio (A (L)/A (S)) and delta(13)C, suggesting that trees with more leaf area for a given sapwood area experienced a stomatal limitation on carbon gain. Nonetheless, these changes had no significant effects on leaf specific conductivities of suppressed versus released trees of either species, but leaf specific root conductance was significantly lower in released Douglas-fir.