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.     

Influence of ammonium and nitrate supply on growth, dry matter partitioning, N uptake and photosynthetic capacity of Pinus radiata seedlings

Growth and physiological responses of Pinus radiata D. Don seedlings to a combination of N supply regimes (low N = 1.78 mol m⁻³, high N = 7.14 mol m⁻³) and ammonium:nitrate ratios (80:20, 50:50 and 20:80; molar basis) were assessed in a hydroponic experiment run over the course of 105 days. Highly significant (P < 0.001) increases in seedling diameter, height, leaf area and dry mass occurred at lower ammonium:nitrate ratios and were two to fourfold greater than the non-significant (for diameter) to marginally significant (P < 0.05 for other dimensions) increases in these dimensions that occurred with greater N supply. Increases in N supply resulted in a highly significant (P < 0.001) reduction in biomass partitioning to roots and highly significant (P < 0.001) increases in allocation to foliage. The ammonium:nitrate ratio was not found to significantly change biomass partitioning to either foliage, stems or roots. Ammonium and nitrate uptake was significantly influenced by N supply and N form and conformed to ammonium and nitrate concentrations in nutrient solution. Uptake rates of ammonium were twice those of nitrate at comparable concentrations suggesting that P. radiata is in the lower end of the ratio of uptake of ammonium to nitrate reported for conifers (range from 2 to 20 mol mol⁻¹). Despite this, plants growing in high ammonium:nitrate ratios were smaller, exhibited luxurious N consumption and lower N use efficiency. Differences in productivity among treatments were partially explained by greater rates of light-saturated photosynthesis associated with nitrate nutrition.     

Stomatal conductance and leaf water potential responses to hydraulic conductance variation in <Emphasis Type="Italic">Pinus pinaster</Emphasis> seedlings

In this study, tree hydraulic conductance (K _tree) was experimentally manipulated to study effects on short-term regulation of stomatal conductance (g _s), net photosynthesis (A) and bulk leaf water potential (Ψ_leaf) in well watered 5–6 years old and 1.2 m tall maritime pine seedlings (Pinus pinaster Ait.). K _tree was decreased by notching the stem and increased by progressively excising the root system and stem. Gas exchange was measured in a chamber at constant irradiance, vapour pressure deficit, leaf temperature and ambient CO₂ concentration. As expected, we found a strong and positive relationship between g _s and K _tree (r = 0.92, P = 0.0001) and between A and K _tree (r = 0.9, P = 0.0001). In contrast, however, we found that the response of Ψ_leaf to K _tree depended on the direction of change in K _tree: increases in K _tree caused Ψ_leaf to decrease from around −1.0 to −0.6 MPa, but reductions in K _tree were accompanied by homeostasis in Ψ_leaf (at −1 MPa). Both of these observations could be explained by an adaptative feedback loop between g _s and Ψ_leaf, with Ψ_leaf prevented from declining below the cavitation threshold by stomatal closure. Our results are consistent with the hypothesis that the observed stomatal responses were mediated by leaf water status, but they also suggest that the stomatal sensitivity to water status increased dramatically as Ψ_leaf approached −1 MPa.     

Transpiration and whole-tree conductance in ponderosa pine trees of different heights

Changes in leaf physiology with tree age and size could alter forest growth, water yield, and carbon fluxes. We measured tree water flux (Q) for 14 ponderosa pine trees in two size classes (12 m tall and ∼40 years old, and 36 m tall and ∼ 290 years old) to determine if transpiration (E) and whole-tree conductance (g _t) differed between the two sizes of trees. For both size classes, E was approximately equal to Q measured 2 m above the ground: Q was most highly correlated with current, not lagged, water vapor pressure deficit, and night Q was <12% of total daily flux. E for days 165–195 and 240–260 averaged 0.97 mmol m^–2 (leaf area, projected) s^–1 for the 12-m trees and 0.57 mmol m^–2 (leaf area) s^–1 for the 36-m trees. When photosynthetically active radiation (I _P) exceeded the light saturation for photosynthesis in ponderosa pine (900 μmol m^–2 (ground) s^–1), differences in E were more pronounced: 2.4 mmol m^–2 (leaf area) s^–1 for the 12-m trees and 1.2 mmol m^–2 s^–1 for the 36-m trees, yielding g _t of 140 mmol m^–2 (leaf area) s^–1 for the 12-m trees and 72 mmol m^–2 s^–1 for the 36-m trees. Extrapolated to forests with leaf area index =1, the 36-m trees would transpire 117 mm between 1 June and 31 August compared to 170 mm for the 12-m trees, a difference of 15% of average annual precipitation. Lower g _t in the taller trees also likely lowers photosynthesis during the growing season. Received: 19 April 1999 / Accepted: 23 March 2000     

Shifts in plant nutrient use strategies under secondary forest succession

In evergreen broad-leaved forests (EBLFs) in Tiantong National Forest Park, Eastern China, we studied the soil chemistry and plant leaf nutrient concentration along a chronosequence of secondary forest succession. Soil total N, P and leaf N, P concentration of the most abundant plant species increased with forest succession. We further examined leaf lifespan, leaf nutrient characteristics and root–shoot attributes of Pinus massoniana Lamb, the early-successional species, Schima superba Gardn. et Champ, the mid-successional species, and Castanopsis fargesii Franch, the late-successional species. These species showed both intraspecific and interspecific variability along succession. Leaf N concentration of the three dominant species increased while N resorption tended to decrease with succession; leaf P and P resorption didn’t show a consistent trend along forest succession. Compared with the other two species, C. fargesii had the shortest leaf lifespan, largest decay rate and the highest taproot diameter to shoot base diameter ratio while P. massoniana had the highest root–shoot biomass ratio and taproot length to shoot height ratio. Overall, P. massoniana used ‘conservative consumption’ nutrient use strategy in the infertile soil conditions while C. fargesii took up nutrients in the way of ‘resource spending’ when nutrient supply increased. The attributes of S. superba were intermediate between the other two species, which may contribute to its coexistence with other species in a wide range of soil conditions.     

Effects of N addition rates on the productivity ofPicea Sitchensis,Thuja plicata,andTsuga heterophylla seedlings

Seedlings ofPicea sitchensis, Thuja plicata andTsuga heterophylla were supplied N hydroponically at one of four exponentially increasing rates of addition (0.09, 0.07, 0.05, or 0.025 gN^-1 day^-1) for up to 3 months in a naturally illuminated glasshouse. Relative growth rates (RGR) were analyzed as a function of N uptake, the allocation of assimilated N to foliage (LNFR), foliar N concentrations (N_la) and met assimilation rates (NAR), which were combined to estimate N productivity (RGR per unit whole-plant N concentration). Nitrogen accumulation, biomass and N partitioning and RGR and its components varied with species in response to the different N regimes.T. heterophylla had the lowest maximum wholeplant N concentrations (wpN) and specific absorption rates for N and exhibited the least plasticity in root: shoot ratios as wpN increased from 11–21 mg g^-1. In all species, RGR increased linearly with wpN, while LNFR increased curvilinearly. Foliar N (N_la) increased linearly with wpN and NAR increased linearly with N_la. The RGRs ofT. heterophylla were highest at wpNs up to 18 mg g^-1, a result of higher foliar N use efficiencies (NAR/N_la). However, RGR increased more with wpN inT. plicata andP. sitchensis. Although LNFR increased with wpN in all species, foliar N use efficiency declined, possibly due to an increased partitioning of foliar soluble N to non-photosynthetic compounds. Thus, in each species, N productivity did not increase above intermediate levels of wpN: 14 mg g^-1 inT. heterophylla, 16 mg g^-1 inP. sitchensis and 17 mg g^-1 inT. plicata.     

Why does photosynthesis decrease with needle age in Pinus pinaster?

Rates of photosynthesis vary with foliage age and typically decline from full-leaf expansion until senescence occurs. This age-related decline in photosynthesis is especially important in species that retain foliage for several years, yet it is not known whether the internal conductance to CO₂ movement (g _i) plays any role. More generally, g _i has been measured in only a few conifers and has never been measured in leaves or needles older than 1 year. The effect of ageing on g _i was investigated in Pinus pinaster, a species that retains needle for 4 or more years. Measurements were made in autumn when trees were not water limited and after leaf expansion was complete. Rates of net photosynthesis decreased with needle age, from 8 μmol m⁻² s⁻¹ in fully expanded current-year needles to 4.4 μmol m⁻² s⁻¹ in 3-year-old needles. The relative limitation due to internal conductance (0.24–0.35 out of 1) was in all cases larger than that due to stomatal conductance (0.13–0.19 out of 1). Internal conductance and stomatal conductance approximately scaled with rates of photosynthesis. Hence, there was no difference among year-classes in the relative limitations posed by internal and stomatal conductance or evidence that they cause the age-related decline in photosynthesis. There was little evidence that the age-related decline in photosynthesis was due to decreases in contents of N or Rubisco. The decrease in rates of photosynthesis from current-year to older needles was instead related to a twofold decrease in rates of photosynthesis per unit nitrogen and V _cmax/Rubisco (i.e., in vivo specific activity).     

Nutrient status in needles of Norway spruce and Scots pine following harvesting of logging residues

Nutrient concentrations in current and 1-year old needles from two Picea abies (L.) Karst and two Pinus sylvestris L. stands in Sweden were determined 8–10, 16–18 and 22–24 years after clear-felling and experimental manipulation of harvesting intensity. On all sites, three levels of harvest intensity had been applied in a randomized block design (n=4); (i) conventional stem-only harvesting, where all logging residues (i.e. tops, branches and needles) were evenly distributed on the ground, (ii) harvesting all above-ground tree parts except needles and (iii) above-ground whole-tree harvesting (no residues left on site). At stand age 8–10 years, nitrogen concentrations in the current year needles in plots where all residues or needles only were retained were higher than in whole-tree harvested plots, whereas concentrations of K, Ca and Mg were lower. The latter response was interpreted as a dilution effect. P:N, K:N, Ca:N, Mg:N, Mn:N and Zn:N were in general higher after whole-tree harvesting treatments than after the treatments where all residues or only needles had been left on site. At stand age 16–18 years, no significant differences in nitrogen concentrations were observed between treatments, but the levels of Ca, Mg and Mn in both current and 1-year-old needles were lower after whole-tree harvesting than after the treatments where logging residues remained on site. By contrast, potassium levels in the foliage were highest in treatments where only the needles were left on site, whereas the lowest levels were observed for treatments where all residues was left. At stand age 22–24 years, the treatment effects had diminished, except for the effects on Ca and K on the southern Norway spruce stand. It is concluded that the nutrient release from logging residues enhances nutrient uptake in trees of the succeeding forest generation, but this effect does not occur simultaneously for all elements. This revised version was published online in June 2006 with corrections to the Cover Date.     

Comparative study of leaf carbon gain in saplings ofThujopsis dolabrata var.hondai andQuercus mongolica var.grosseserrata in a cool-temperate deciduous forest

Seasonal courses of leaf CO₂ gas exchange in a growing season were examined in saplings ofThujopsis dolabrata var.hondai andQuercus mongolica var.grosseserrata in a cool temperate deciduous forest. Between the two tree species there were no large differences in the light compensation point of leaf photosynthesis, except for the season of new leaf expansion. However, light-saturated rates of net photosynthesis were obviously high inT. dolabrata var.hondai. EvergreenT. dolabrata var.hondai saplings had large photosynthetic production in two seasons, before the emergence of new foliage and after foliage fall of the overstory deciduous trees, because of the significantly high solar radiant energy penetrating under the forest canopy during the seasons. Saplings of deciduousQ. mongolica var.grosseserrata were heavily shaded throughout the growing season by foliage of the overstory trees, which resulted in a low daily surplus production. The annual surplus production of leaves in the growing season was estimated to be 2300 mmol CO₂ m⁻² inT. dolabrata var.hondai and −100 mmol CO₂ m⁻², slightly negative, inQ. mongolica var.grosseserrata. These results supported the high survivability ofT. dolabrata var.hondai saplings and the high mortality ofQ. mongolica var.grosseserrata in the deciduous forest.     

Regulation of water flux through trunks, branches, and leaves in trees of a lowland tropical forest

We studied regulation of whole-tree water use in individuals of five diverse canopy tree species growing in a Panamanian seasonal forest. A construction crane equipped with a gondola was used to access the upper crowns and points along the branches and trunks of the study trees for making concurrent measurements of sap flow at the whole-tree and branch levels, and vapor phase conductances and water status at the leaf level. These measurements were integrated to assess physiological regulation of water use from the whole-tree to the single-leaf scale. Whole-tree water use ranged from 379 kg day⁻¹ in a 35 m-tall Anacardium excelsum tree to 46 kg day⁻¹ in an 18 m-tall Cecropia longipes tree. The dependence of whole-tree and branch sap velocity and sap flow on sapwood area was essentially identical in the five trees studied. However, large differences in transpiration per unit leaf area (E) among individuals and among branches on the same individual were observed. These differences were substantially reduced when E was normalized by the corresponding branch leaf area:sapwood area ratio (LA/SA). Variation in stomatal conductance (g _s) and crown conductance (g _c), a total vapor phase conductance that includes stomatal and boundary layer components, was closely associated with variation in the leaf area-specific total hydraulic conductance of the soil/leaf pathway (G _t). Vapor phase conductance in all five trees responded similarly to variation in G _t. Large diurnal variations in G _t were associated with diurnal variation in exchange of water between the transpiration stream and internal stem storage compartments. Differences in stomatal regulation of transpiration on a leaf area basis appeared to be governed largely by tree size and hydraulic architectural features rather than physiological differences in the responsiveness of stomata. We suggest that reliance on measurements gathered at a single scale or inadequate range of scale may result in misleading conclusions concerning physiological differences in regulation of transpiration. Received: 1 October 1997 / Accepted: 6 March 1998     

Growth and uptake of N,P, K and B by Pinus radiata D. Don in response to applications of borax

Leader dieback associated with B deficiency in P. radiata D. Don plantations was treated with borax applied at rates of 50, 100 and 150 kg ha⁻¹. This initially increased B in foliage from 5 to 40, 80 and 110 μg g⁻¹ respectively, and was followed by a rapid decline and stabilisation at around 25 μg g⁻¹ for the duration of the study. Annual fluctuations in foliage B levels were strongly correlated with rainfall during the preceding spring and summer. Uptake of N, P and K increased as a result of applied B and comparison of the distribution of these nutrients in crowns of fertilized and unfertilised trees six years after application indicated continued uptake of these nutrients probably as a result of improved root growth due to B. Foliage concentrations of B like N, P and K, increased in young needles towards the upper crown and this, together with a decline in needle concentrations of B as foliage aged, indicated some redistribution of B from older to new foliage. A limit of 5 μg g⁻¹ was found below which little redistribution seems to occur. Application of B prevented further leader dieback, improved apical dominance and height growth and increased volume production by 25 m³ ha⁻¹ at age 8 years. Differences between application rates of B were not significant in terms of growth.     

Ontogenetic differentiation between Mediterranean and Eurasian pines (sect. Pinus) at the seedling stage

Heteroblastic development in pine seedlings includes extreme morphological changes with still unclear adaptive and evolutionary significance. In particular, Mediterranean and Eurasian pines (section Pinus) living in the Mediterranean basin seem to follow quite distinct developmental trajectories at the seedling stage. Aiming to confirm this ontogenetic differentiation we performed a nursery experiment with seedlings of five Mediterranean pines (Pinus pinaster, P. brutia, P. halepensis, P. pinea and P. canariensis) and three Eurasian pines (P. sylvestris, P. uncinata, and P. nigra subsp. salzmannii), also including P. radiata as an outgroup. After destructive analyses at two harvest times (9 and 32 weeks), we found sharp differentiation between Mediterranean and Eurasian pines in a combination of traits linked to shoot heteroblasty. In particular, Mediterranean pines showed a marked delay in the proportion of adult needles to total needles in the shoot compared to Eurasian species, especially at the second harvest. However, two Mediterranean pines, P. pinaster and P. brutia showed a slightly higher proportion of secondary needles, and a higher rate of budset at a more advanced stage (68 weeks) compared to the other three Mediterranean species. Meaningfully, the outgroup P. radiata was the only species combining a high proportion of adult foliage since the first harvest with a delayed formation of the first terminal bud. We discussed the adaptive significance of these findings at the light of species’ climatic niches and life histories.     

Estimates of biomass and primary productivity in a high-altitude maple forest of the west central Himalayas

The paper describes the biomass and productivity of maple (Acer cappadocicum) forest occurring at an altitude of 2,750 m in the west central Himalayas. Total vegetation biomass was 308.3 t ha⁻¹, of which the tree layer contributed the most, followed by herbs and shrubs. The seasonal forest-floor litter mass varied between 5.4 t ha⁻¹ (in rainy season) and 6.6 t ha⁻¹ (in winter season). The annual litter fall was 6.2 t ha⁻¹, of which leaf litter contributed the largest part (59% of the total litter fall). Net primary productivity of total vegetation was 19.5 t ha⁻¹ year⁻¹. The production efficiency of leaves (net primary productivity/leaf mass) was markedly higher (2.9 g g⁻¹ foliage mass year⁻¹) than those of the low-altitude forests of the region.     

Alleviation of Ultraviolet-B Radiation-Induced Growth Inhibition of Green Gram by Triadimefon

Supplementary UV-B (12.2 kJ m⁻² d⁻¹ UV-B_BE) provided to Vigna radiata for 2 h d⁻¹ suppressed the length of root, shoot and whole plants, number of leaves, total leaf area, leaf area index, specific leaf mass, fresh and dry mass of leaves and shoot, relative growth rate and net productivity. In unstressed green gram plants (10 kJ m⁻² d⁻¹ UV-B_BE), triadimefon (TRIAD) (20 mg dm⁻³) enhanced growth in all parameters over control. The growth promoting effect of TRIAD enabled the UV-B impacted plants to overcome the growth inhibitions to varying degrees indicating its protective potential against UV-B stress. This revised version was published online in July 2006 with corrections to the Cover Date.     

Clonal micropropagation of <Emphasis Type="Italic">Crataeva adansonii</Emphasis> (DC.) Prodr.: A multipurpose tree

Summary A method of micropropagation through multiple shoot formation from axillary buds of mature tree and rootstock growths of Crataeva adansonii (DC.) Prodr. (a multipurpose tree) has been developed. Factors affecting multiplication rate included season, age of explant source, explant type, type of bud, position of bud on the foliage twig, type of medium, various additives, and explant implantation on the medium. The maximum number of buds was produced from the sixth to 10th axillary buds taken from foliage twigs of 40–50-d-old rootstock growth in the months of October to December. At this time of the year the contamination was minimum. Optimum response was recorded on Murashige and Skoog (MS) medium supplemented with 6-benzyladenine (BA; 3 mgl⁻¹, 13.3μM) and 1-naphthaleneacetic acid (NAA; 0.05 mgl⁻¹, 0.27 μM) after 21 d of culture. Shoot buds were further multiplied and maintained on BA (1–1.5 mgl⁻¹, 4.4–6.7 μM) while shoots elongated on BA (0.1–0.5 mgl⁻¹, 0.44–2.2 μM) supplemented medium. The number of shoots was further multiplied by using nodal segments of in vitro-regenerated shoots as microcuttings and repeated subculturing of stumps after excising the microshoots. In vitro rooting on growth regulator-free MS medium was possible with 70% of microshoots after 4 wk. From one nodal segment 150 plantlets were produced within 14 wk.     

Carbon and nitrogen pools in a mangrove stand of <Emphasis Type="Italic">Kandelia obovata</Emphasis> (S., L.) Yong: vertical distribution in the soil–vegetation system

Attempts were made to quantify the carbon and nitrogen pools in a monospecific and pioneer mangrove stand of Kandelia obovata Sheue, Liu & Yong, Okinawa Island, Japan. The leaf C and N concentrations on a leaf area basis decreased with increasing PPFD (Photosysthetic Photon Flux Density). The total C and N stocks in foliage were estimated as 3.55 Mg ha^–1 and 0.105 Mg ha^–1, respectively. The bark (45.6–48.6% for C and 0.564–0.842% for N) contained significantly higher amount of C (P < 0.05) and N (P < 0.01) than wood (46.2–47.8% for C and 0.347–0.914% N). The total C stock of stem was 23.2 Mg ha^–1 in wood and 8.33 Mg ha^–1 in bark, and the total N stock was 0.222 Mg ha^–1 in wood and 0.116 Mg ha^–1 in bark. The root wood (37.1–45.0%) contained significantly higher amount of C than root bark (35.4–40.7%) (P < 0.01). The total C stock of root was 14.2 Mg ha^–1 in wood and 12.6 Mg ha^–1 in bark, and the total N stock of root was 0.157 Mg ha^–1 in wood and 0.155 Mg ha^–1 in bark. The soil organic C and total N stocks within 1 m soil depth were estimated as 57.3 Mg ha^–1 and 2.73 Mg ha^–1, respectively. The C pool in aboveground biomass (35.1 Mg ha^–1) was 1.3 times as large as that in belowground biomass (26.9 Mg ha^–1). However, the soil organic C pool (57.3 Mg ha^–1) was similar to the total C pool (62.0 Mg ha^–1) of vegetation, indicating that the mangrove stored a large part of production in the soil. About 50% of the C was in the soil. The N pool in aboveground biomass (0.442 Mg ha^–1) was 1.4 times as large as that in belowground biomass (0.312 Mg ha^–1). The soil N stock was 3.3 times as large as the biomass N stock (0.754 Mg ha^–1).     

Different nutrient use strategies of expansive grasses <Emphasis Type="Italic">Calamagrostis epigejos</Emphasis> and <Emphasis Type="Italic">Arrhenatherum elatius</Emphasis>

Enhanced nitrogen (N) levels accelerate expansion of Calamagrostis epigejos and Arrhenatherum elatius, highly aggressive expanders displacing original dry acidophilous grassland vegetation in the Podyjí National Park (Czech Republic). We compared the capability of Calamagrostis and Arrhenatherum under control and N enhanced treatments to (i) accumulate N and phosphorus (P) in plant tissues, (ii) remove N and P from above-ground biomass during senescence and (iii) release N and P from plant material during decomposition of fresh formed litter. In control treatment, significantly higher amounts of total biomass and fresh aboveground litter were observed in Calamagrostis than in Arrhenatherum. Contrariwise, nutrient concentrations were significantly higher (11.6–14.3 mg N g⁻¹ and 2.3 mg P g⁻¹) in Arrhenatherum peak aboveground biomass than in Calamagrostis (8.4–10.3 mg N g⁻¹ and 1.6–1.7 mg P g⁻¹). Substantial differences between species were found in resorption of nutrients, mainly P, at the ends of growing seasons. While P concentrations in Arrhenatherum fresh litter were twice and three times higher (1.6–2.5 mg P g⁻¹) than in Calamagrostis (0.7–0.8 mg P g⁻¹), N concentrations were nearly doubled in Arrhenatherum (13.1–15.6 mg N g⁻¹) in comparison with Calamagrostis (7.4–8.7 mg N g⁻¹). Thus, the nutrients (N and mainly P) were retranslocated from the aboveground biomass of Calamagrostis probably more effectively in comparison with Arrhenatherum at the end of the growing season. On the other hand, Arrhenatherum litter was decomposed faster and consequently nutrient release (mainly N and P) was higher in comparison with Calamagrostis which pointed to different growth and nutrient use strategies of studied grass species.     

Phenotypic plasticity and biomass allocation pattern in three <Emphasis Type="Italic">Dryopteris</Emphasis> (Dryopteridaceae) species on an experimental light-availability gradient

We were interested in whether the contrasting regional distribution patterns of three congeneric, frequently co-occurring fern species (Dryopteris carthusiana, D. dilatata and D. expansa) could be explained by differential biomass allocation strategies and different phenotypic plasticities to light availability. The morphology and habitat preference of these ferns are known to be very similar, but in Estonia, their frequencies of occurrence differ sharply––Dryopteris carthusiana is common, D. expansa grows in scattered localities, and D. dilatata is rare. We grew the species under different levels of illumination (100, 50, 25 and 10% of full daylight) in an experimental garden to compare their autecological responses to shading. After one growing season there were clear interspecific differences in total plant biomass accumulation––D. carthusiana > D. expansa > D. dilatata––indicating the possible competitive inferiority of the latter at the young sporophyte stage. D. expansa was the least shade-tolerant, with biomass decreasing sharply under less than 50% illumination; D. dilatata was the most shade-tolerant, with similar growth at all illumination levels. In relative biomass allocation patterns, the most notable differences among species were in the relative shares of biomass stored in rhizomes. In D. carthusiana and D. expansa this share was nearly constant and independent of the illumination conditions. D. dilatata allocated very little biomass into rhizome in deep shade, but was able to increase this share more than twofold in 50% light. Dryopteris dilatata was clearly shown to be morphologically the most plastic of the three. In four traits––rhizome mass, frond:below-ground biomass ratio, stipe length and specific leaf area––its degree of ontogenetic plasticity to light was significantly higher than that of D. expansa and D. carthusiana. While the general performance (biomass production) of species in the experiment coincided with that observed in nature, the results of plasticity estimation were somewhat surprising––it is difficult to explain the inferior performance of a species (D. dilatata) through high morphological plasticity. Probably, the species is rare either because of certain climatic restrictions, or because it is presently expanding its distribution and is in the phase of invading Estonian understory communities.     

Gas-exchange and sap flow measurements of Salix viminalis trees in short-rotation forest

Simultaneous field measurements of transpiration and sap flow were performed on short-rotation Salix viminalis trees ranging in diameter from 1.5 to 3.5 cm (2-year-old shoots on 8-year-old stumps). Transpiration was measured using an open-top ventilated chamber enclosing the whole foliage of a tree. Sap flow was measured using a tree-trunk heat balance (THB) technique with a constant temperature difference and variable heat input. Both the instantaneous and daily values of water flux measured by the two absolute techniques agreed well with a difference of up to about 5%. In July, the hourly transpiration reached a maximum of about 0.2 kg m^–2 (leaf area) or 0.45 kg tree^–1, whereas maximum daily integrals reached 4 kg tree^–1. The response of sap flow rate to abrupt flux change when inducing emboli by cutting-off the stem was very rapid: the registered signal dropped by 85% within 10 min for a specimen with a projected leaf area of 2 m². For S. viminalis trees, transpiration was linearly correlated with stem cross-sectional area and with leaf area.     

Altitudinal Change in LAI and Stand Leaf Biomass in Tropical Montane Forests: a Transect Study in Ecuador and a Pan-Tropical Meta-Analysis

Abstract Leaf area index (LAI) is a key parameter controlling plant productivity and biogeochemical fluxes between vegetation and the atmosphere. Tropical forests are thought to have comparably high LAIs; however, precise data are scarce and environmental controls of leaf area in tropical forests are not understood. We studied LAI and stand leaf biomass by optical and leaf mass-related approaches in five tropical montane forests along an elevational transect (1,050–3,060 m a.s.l.) in South Ecuador, and conducted a meta-analysis of LAI and leaf biomass data from tropical montane forests around the globe. Study aims were (1) to assess the applicability of indirect and direct approaches of LAI determination in tropical montane forests, (2) to analyze elevation effects on leaf area, leaf mass, SLA, and leaf lifespan, and (3) to assess the possible consequences of leaf area change with elevation for montane forest productivity. Indirect optical methods of LAI determination appeared to be less reliable in the complex canopies than direct leaf mass-related approaches based on litter trapping and a thorough analysis of leaf lifespan. LAI decreased by 40–60% between 1,000 and 3,000 m in the Ecuador transect and also in the pan-tropical data set. This decrease indicates that canopy carbon gain, that is, carbon source strength, decreases with elevation in tropical montane forests. Average SLA decreased from 88 to 61 cm² g⁻¹ whereas leaf lifespan increased from 16 to 25 mo between 1,050 and 3,060 m in the Ecuador transect. In contrast, stand leaf biomass was much less influenced by elevation. We conclude that elevation has a large influence not only on the leaf traits of trees but also on the LAI of tropical montane forests with soil N (nitrogen) supply presumably being the main controlling factor.