Influence of several tree traits on rainfall partitioning in temperate and boreal forests: a review

? In forests, rainfall is partitioned into intercepted water (IW), throughfall (TF) and stemflow (SF). We reviewed the majority of published works comparing water flows among tree species in temperate and boreal forests to test the effect of several tree traits on water flows.

? We hypothesized that water flows differed between evergreen and deciduous species, and according to successional status and bark roughness. We also investigated that water flows would be explained by stand variables such as basal area.

? Linear mixed models fitted on reviewed data showed that evergreens had a lower TF than deciduous trees (decrease of 13.9% of total precipitation year-round and 8.4% over the growing period). Similar results were found for conifers compared to broadleaves. TF also declined with the successional status from pioneer to late-successional tree species. SF decreased with bark roughness from smoother to rougher bark. Evergreens had water flows that were dependent on age of the stand, especially for TF which increased by 15.6% of total precipitation from young to adult forests.

? The large scale of TF differences between tree genera together with specific transpiration amounts and rooting features highlighted in other studies should result in significant differences in soil water content among tree species. This may have consequences on stand fitness and growth, and understory vegetation.

     

Hybridization between Quercus robur and Q. petraea in a mixed oak stand in Denmark

? Hybridization and mating pattern between Quercus robur and Q. petraea was studied in a 5.8 ha mixed forest stand in Jutland, Denmark which comprises in total 135 Quercus robur and 230 Q. petraea trees. Classification of the oak trees into species was performed using canonical discriminant analysis of a range of leaf morphological traits. Adult trees (365) and offspring (582) were genotyped with eight microsatellite markers. Seedlings were sampled in 2003 and acorns were collected in 2004.

? Mating patterns of Q. robur and Q. petraea are expected to be different in the northern range of the distribution area and a larger hybridization rate is expected. It is further expected, that pollination from outside sources will be relatively less in small fragmented forest management systems compared to large scale oak forest. The conclusions should be verified through repeated year to year analysis of the mating pattern.

? Phenological studies revealed that there was no difference in flowering time between species. Data for the adult trees revealed no significant departures from Hardy-Weinberg proportions and there was weak, but significant spatial genetic structure, which supports the idea that the stand is of natural origin. Spatial genetic structure in the first distance class is stronger for Q. petraea. The genetic composition of the offspring was remarkably consistent from year to year. Paternity analysis revealed that, on average, 85% pollination came from fathers within the stand. The direction of the pollen flow varied from year to year. Inter-specific hybridization was high and ranged from 15–17% and from 48–55% for Q. petraea and Q. robur mothers respectively. Paternity analysis revealed that the population was basically outcrossing and only 3.7% of the analysed progeny were the product of selfing. Over the two years of study, approximately 200 trees contributed to the paternity of the next generations.

? The study confirms earlier studies with a greater tendency for Q. robur mothers to produce hybrid seeds than Q. petraea mothers. The rate of hybridization is higher in this Danish stand than in comparable studies elsewhere in Europe. Gene flow from outside sources are relatively low.

     

Productivity of temperate broad-leaved forest stands differing in tree species diversity

? Understanding the effects of tree species diversity on biomass and production of forests is fundamental for carbon sequestration strategies, particularly in the perspective of the current climate change. However, the diversity-productivity relationship in old-growth forests is not well understood.

? We quantified biomass and above-ground production in nine forest stands with increasing tree species diversity from monocultures of beech to stands consisting of up to five deciduous tree species (Fagus sylvatica, Fraxinus excelsior, Tilia spp., Carpinus betulus, Acer spp.) to examine (a) if mixed stands are more productive than monospecific stands, (b) how tree species differ in the productivity of stem wood, leaves and fruits, and (c) if beech productivity increases with tree diversity due to lower intraspecific competition and complementary resource use.

? Total above-ground biomass and wood production decreased with increasing tree species diversity. In Fagus and Fraxinus, the basal area-related wood productivity exceeded those of the co-occurring tree species, while Tilia had the highest leaf productivity. Fagus trees showed no elevated production per basal area in the mixed stands.

? We found no evidence of complementary resource use associated with biomass production. We conclude that above-ground productivity of old-growth temperate deciduous forests depend more on tree species-specific traits than on tree diversity itself.

     

Characteristics of a 20-year-old evergreen broad-leaved forest restocked by natural regeneration after clearcut-burning

? To elucidate the application of natural regeneration to the restocking of evergreen broad-leaved forests in the subtropics, the characteristics of a 20-year-old evergreen broad-leaved forest restocked by natural regeneration after clearcut-burning were studied in Okinawa, Japan. Within a 0.87 ha clearcut area with four 10 m × 10 m sampling plots, two burned and two unburned ones, the tree composition, tree species diversity and vegetation changes were investigated.

? The results showed that the species diversity, basal area and density of woody stems ? 1.0 m in height differed significantly among phanerophyte types, while they were not significantly different between the burned and unburned treatments. A vegetation census also revealed no obvious differences between the treatments.

? The primary dominant species, Castanopsis sieboldii, continued to dominate the secondary forest with a broad height distribution.

? The structural complexity and high tree species diversity of the regenerating forest after clearcut-burning provides no evidence of degeneration. We can predict that the regeneration forest may gradually develop into stands similar to pre-clearcut primary forest, and that natural regeneration may restore the high tree species diversity of the evergreen broad-leaved forests in Okinawa.

     

Photosynthetic temperature adaptation of Pinus cembra within the timberline ecotone of the Central Austrian Alps

? Temperature is suggested to determine the upper limit of tree life. Therefore, future climate warming may be of importance for tree distribution within the European Alps, where low temperatures limit carbon metabolism.

? We focused on the effects of air and soil temperature on net photosynthesis (P _n) of Pinus cembra an evergreen climax species of the timberline ecotone of the Central Austrian Alps. Light response and temperature response curves were estimated along an altitudinal gradient ranging from the forest limit up to the krummholz limit in both summer and fall.

? In general, P _n was significantly lower in fall as compared to summer. Nevertheless, independent from season mean P _n values tended to increase with elevation and were positively correlated with root zone temperatures. The specific leaf area by contrast declined with increasing elevation. Furthermore, the temperature optimum of net photosynthesis declined with increasing elevation and was positively correlated with the mean maximum air temperature of the 10 days prior the date of measurement.

? Thus, our findings appear to reflect a long-term adaptation of the photosynthetic apparatus of Pinus cembra to the general temperature conditions with respect to elevation combined with a short term acclimation to the prevailing temperature regime.

     

Cold hardiness and transplant response of Juglans nigra seedlings subjected to alternative storage regimes

? Effects of overwinter storage regimes on seedling cold hardiness and physiological vigor are relatively unexplored, particularly for temperate deciduous forest tree species.

? We evaluated influence of storage duration (0, 66, 119, or 175 d) on electrolyte leakage of stem and root collar tissues following exposure to a series of freeze-test temperatures in black walnut (Juglans nigra L.) seedlings sampled from cold (3 °C) or freezer (?2 °C) storage. Seedlings were subsequently transplanted into a controlled growth chamber environment for two months.

? Regardless of storage temperature, mean LT₅₀ was lowest for seedlings stored for 66 d (≤ ?34 °C) and increased dramatically after 119 d (≥ ?13 °C).

? Root collar tissue had lower LT₅₀ than stem tissue after 119 d for cold-stored seedlings, reflecting importance of evaluative tissue type. Days to bud break shortened with increasing storage duration up to 119 d and stabilized thereafter for both storage regimes. Root growth potential was maximized after 119 d of storage, and subsequently declined for cold-stored seedlings. Height growth increased following storage, regardless of duration.

? To promote stress resistance and transplant growth response, we recommend that black walnut seedlings from this genetic source be outplanted after approximately 66–119 d of storage.

     

Temporal trends in the foliar nutritional status of the French, Walloon and Luxembourg broad-leaved plots of forest monitoring

? Two decades after the launching of the monitoring program of forest ecosystems in Europe (ICP forests), a unique data set is now available regarding the foliar nutritional status of the main broad-leaved species growing in much-diversified sites (soil, climate).

? This study focuses on the foliar concentration time series (1993 to 2005) of the French (RENECOFOR), Walloon and Luxemburg broad-leaved plots. The aim is to show long-term trends while taking the inter-plot and inter-annual variability into account.

? Two kinds of statistical processing were used to analyze the data on foliar chemistry: principal component analysis (PCAs) and linear mixed models. In general, the main temporal trends reveal a decrease in the foliar P concentration and a decrease in the foliar Ca concentration, except for European beech on acid soils.

? These temporal patterns confirm the trends previously observed and could be explained by the joint actions of several processes that influence tree nutrition in the long term: tree age, N and S deposition, harvesting and climate.

     

Winter variation in physiological status of cold stored and freshly lifted semi-evergreen Quercus nigra seedlings

? Water oak (Quercus nigra L.) is a tardily deciduous species commonly planted in afforestation projects in the Lower Mississippi River Alluvial Valley, USA. Field performance is often marked by low survival rates and top dieback, which may be associated with poor physiological quality of planting stock.

? We investigated physiological status of cold stored (2–4 °C; CS) and freshly lifted (FL) seedlings during the period between lifting and planting (December — February). In mid-February, seedlings were transplanted into a controlled greenhouse environment for 90 d to evaluate post-transplant growth performance.

? Net photosynthetic rates were positive until late January (generally greater in CS seedlings) and became negative thereafter. FL seedlings generally had lower LT₅₀ values from freeze-induced electrolyte leakage (FIEL), reflecting greater cold hardiness. FIEL of foliage provided the best indicator of physiological status, though terminal buds may serve as a suitable substitute. All seedlings experienced top dieback following transplant; CS seedlings had less relative root-collar diameter, height, and root volume increments.

? Cold storing seedlings did not appear to prolong dormancy, increase stress resistance, or hold promise as a means to improve outplanting success. Regardless of storage regime, seedlings appeared to be most cold hardy and perhaps stress resistant until late January.

     

Estimation of chlorophyll in Quercus leaves using a portable chlorophyll meter: effects of species and leaf age

? The potential of a chlorophyll meter (CCM-200, Opti-Sciences, Tyngsboro, MA, USA) for the estimation of total leaf chlorophyll contents was evaluated in leaves of different developmental stages for Quercus pyrenaica Willd., Q. faginea Lam., Q. suber L. and Q. ilex L. subsp. ballota (Desf.) Samp.

? For all species and leaf stages, a strong non-linear relationship (P < 0.0001) was found between the chlorophyll content index (CCI) and leaf chlorophyll content per unit area. In all species, the slope of the CCI-chlorophyll content relationship varied during leaf development, suggesting that caution should be exercised when using the CCM-200, since the interpretation of CCI readings should be limited to samples of similar leaf age. Also, the data indicate that the models should also be speciesspecific, owing to differences in the intercept and/or slope of the equations.

? The addition of fresh LMA as a complementary parameter improved the accuracy of chlorophyll estimation in non-species-specific equations to a considerable extent when leaf samples of similar development were considered.

? After due consideration, it may be concluded that the CCM-200 chlorophyll meter is a reliable method for acquiring an estimation of chlorophyll contents in Mediterranean Quercus species, with potential use as an effective tool in forest management.

     

Frost damage in Pinus sylvestris L. stems assessed by chlorophyll fluorescence in cortical bark chlorenchyma

? Several techniques were used to investigate the threshold minimum temperature inducing severe frost damage during winter in Scots pine (Pinus sylvestris L.) needles and stems from southern provenances in the Iberian Peninsula.

? Chlorophyll (Chl) fluorescence, electrolyte leakage (EL), visual scoring (VS), and the normalized difference vegetation index (NDVI) were compared.

? Chl fluorescence, and in particular the maximum potential photosystem II (PSII) efficiency — estimated through the dark-acclimated F_V/F_M ratio-, was found to be a simple, non-destructive indicator of freezing tolerance in needles, as expected.

? Moreover, the existence of a Chl-containing tissue, the cortical bark chlorenchyma, allowed us the use of Chl fluorescence as a new non-destructive indicator of frost damage in live tissues of Pinus sylvestris L. stems.

? Freezing tolerance values of southern provenances are compared with those found by other authors in the northern distribution limit of the species.

     

Responses of photosynthesis to high temperatures in oak saplings assessed by chlorophyll-a fluorescence: inter-specific diversity and temperature-induced plasticity

? The sensitivity of photosynthesis to high temperature was investigated in greenhouse-grown saplings of three Mediterranean (Quercus canariensis Willd., Q. coccifera L. and Q. suber L.) and two temperate European oak species (Q. robur L. and Q. petraea Matt. Liebl.). The sensitivity to high temperature was inferred from F ₀T curves producing the critical temperature (T _c ) at which a rapid rise in ground chlorophyll a fluorescence (F ₀) occurs. T _c is known to acclimate to moderately increased ambient temperature by displaying significant increases.

? We first checked whether acclimation of T _c was paralleled by an increase in optimal temperature for light driven electron flux (T _opt ,), recorded by measuring quantum yield of photochemistry under saturating CO₂ and moderate irradiance. Saplings of Q. suber and Q. canariensis were submitted to four, five days long ambient temperature steps (15, 25, 35 and 15 °C again). T _c was usually largely above 45 °C while T _opt was around 30–35 °C. Both increased with ambient temperature in the two species, and decreased again when temperature returned to the initial values. This recovery was nevertheless not complete after a week at 15 °C.

? A second experiment tested whether the plasticity of T _c was of the same extent in the five species. A significant increase of T _c followed by a decrease was detected in all species. Only small inter-specific differences were detected in T _c , and temperature induced plasticity was similar in the different species: T _c increased by about 4–5 °C while ambient temperature was increased from 15 to 35 °C. Mediterranean and European oaks did not differ in this respect.

? This result does not support the hypothesis that oak species from warmer regions would display a larger plasticity of T _c in response to increasing temperatures.

     

Tree-ring Δ13C reveals the impact of past forest management on water-use efficiency in a Mediterranean oak coppice in Tuscany (Italy)

? The effects of thinning and heavy stand density reduction was investigated in Turkey oak (Quercus cerris L.) forests of central Italy, to evaluate the physiological responses and the growth status of trees that survived a past coppice cut and thinning to convert the stand to high-forest.

? The working hypothesis was that a strong decrease in stand density would cause a decreasing in canopy-intrinsic water-use efficiency (measured as the ratio of CO₂ assimilation to stomatal conductance, A/g), thus an increase in tree-ring carbon isotopic discrimination (Δ¹³C).

? The tree-ring Δ¹³C of the remaining trees (“survivors”) was found to have significantly (P < 0.05) raised between year two and year seven since the coppice stand was thinned (high-forest conversion thinnings). This effect was mostly caused by a large decrease in tree-rings Δ¹³C at control site which was characterized by high density and competition by trees. An increase in survivors tree-rings Δ¹³C probably indicates an improved water availability, possibly induced by a decrease in competition and in stand density or a decrease in the precipitation intercepted by the canopy (i.e., a stronger increase in g over A since a decrease in A is highly unlikely). A change in foliar nitrogen, foliar Δ¹³C and content in chlorophylls was also recorded seven years after thinning.

? Thinnings carried out to convert old abandoned coppices into high-forest stands induce short-term stimulation of Turkey oak growth by increasing light and water availability. We were able to make a detailed reconstruction of the impact of past silvicultural treatment on the stand using a tree-ring wood Δ¹³C time-series.

     

Evidence of altitudinal increase in photosynthetic capacity: gas exchange measurements at ambient and constant CO2 partial pressures

? Because all microclimatic variables change with elevation, it is difficult to compare plant performance and especially photosynthetic capacity at different elevations. Indeed, most previous studies investigated photosynthetic capacity of low- and high-elevation plants using constant temperature, humidity and light but varying CO₂ partial pressures (P _{CO 2}).

? Using gas exchange measurements, we compared here maximum assimilation rates (A _max) at ambient and constant-low-elevation P _{CO 2}for two temperate tree species along an altitudinal gradient (100 to 1600 m) in the Pyrénées mountains.

? Significant differences in A _max were observed between the CO₂ partial pressure treatments for elevations above 600 m, the between-treatment differences increasing with elevation up to 4 μmol m^?2 s^?1. We found an increase in A _max with increasing elevation at constant-low-elevation P _{CO 2} but not at ambient P _{CO 2} for both species. Given a 10% change in P _{CO 2}, a proportionally higher shift in maximum assimilation rate was found for both species.

? Our results showed that high elevation populations had higher photosynthetic capacity and therefore demonstrated that trees coped with extreme environmental conditions by a combination of adaptation (genetic evolution) and of acclimation. Our study also highlighted the importance of using constant CO₂ partial pressure to assess plant adaptation at different elevations.

     

Genetic diversity in Tunisian Crataegus azarolus L. var. aronia L. populations assessed using RAPD markers

? The genetic diversity of nine wild Tunisian Crataegus azarolus var. aronia L. populations from different bioclimates was assessed using RAPD markers.

? Eight selected primers generated a total of 105 bands, 81 of which were polymorphic. Shannon’s index (H′) ranged from 0.222 to 0.278 according to a population with an average of 0.245. The genetic variation within the species (H _SP = 0.423) was relatively low. A high differentiation (G _ST = 0.421) among populations coupled with a low level of gene flow (N _m = 0.472) were observed. The analysis of molecular variance (AMOVA) revealed also significant differentiation among populations (Φ_ST = 0.371), even at a low scale space. The majority of variation occurred within populations (63.31%). The Mantel test performed on genetic (Φ_ST) and geographic distance matrices among population pairs did not reveal an isolation by distance.

? Interpretation of Neighbour-joining tree based on Nei’s and Li’s genetic distance among individuals showed distinct population groupings. The UPGMA dendrogram based on Φ_ST values revealed two population sub-clusters, each including populations from different bioclimates and/or geographic regions.

? The low level of genetic diversity and the high genetic structure of populations resulted from genetic drift caused both by habitat fragmentation and the low size of populations.

? The high differentiation among populations and the similar low level of diversity within populations suggest that in situ conservation should interest all populations. The ex situ conservation should be based on the collection of seeds rather within than among populations because of the maximum of variation was revealed within populations.

     

Leaf water status and stem xylem flux in relation to soil drought in five temperate broad-leaved tree species with contrasting water use strategies

? Five temperate broad-leaved tree species were compared with respect to their water consumption strategies under ample and restricted water supply. We measured synchronously leaf conductance (g _L) in the sun canopy, xylem sap flux (J _s) and leaf water potential (predawn, ψ_pd and noon, ψ_noon) in adult trees in a mixed stand and related them to the fluctuations in vapor pressure deficit (D) and soil moisture.

? Maximum g _L was particularly high in F. excelsior, C. betulus and T. cordata and revealed a higher D sensitivity. ψ_pd remained constantly high in A. pseudoplatanus, C. betulus and F. excelsior, but decreased in T. cordata and F. sylvatica with decreasing soil moisture.

? J _sddecreased linearly with decreasing soil matrix potential in all species except for F. excelsior. Apparent hydraulic conductance in the soil-to-leaf flow path (L _c) was higher in A. pseudoplatanus than in the other species.

? F. sylvatica maintained a low maximum g _L and reduced J _sd markedly upon drought, but faced severe decreases in ψ_pd and ψ_noon. F. excelsior represents an opposite strategy with high maximum g _L and stable ψ_pd.

? The species drought sensitivity increases in the sequence F. excelsior < C. betulus < T. cordata < A. pseudoplatanus < F. sylvatica.

     

Establishment of an agroforestry research network in Victoria,Australia (1983–85)

From 1983 to 1985, a network of agroforestry research trials was established at six localities throughout Victoria. This paper describes experimental designs and establishment procedures used at the trial sites. The overall aim of the research program is to provide definitive information on the costs and benefits of a wide range of agroforestry combinations. Accordingly, trials were established at the following locations to test a total of 93 tree species grown in combination with specific types of agriculture:

1. Carngham (near Ballarat) — sheep grazing andPinus radiata D. Don (radiata pine);
2. Hamilton (several sites) — sheep grazing and mixed tree species;
3. Kyabram — irrigated pastures andEucalyptus grandis W. Hill ex Maiden (flooded gum);
4. Rutherglen — cereal cropping and shelterbelts;
5. Myrtleford — irrigatedMentha spp. (mint) and two clones ofPopulus spp. (poplar); and
6. Neerim South (near Warragul) — cattle grazing and mixed tree species.

Conventional and systematic experimental designs have been used in setting up these trials, which in total constitute the most comprehensive agroforestry research in Australia. This paper describes these trials in detail, and stresses the importance of their continued expert management, so that the full benefits of the work can be realised for rural Victoria.     

Epicormics in 13-year-old Quercus petraea: small effect of provenance and large influence of branches and growth unit limits

? The provenance effect on epicormics is poorly documented and restricted to epicormic shoots. Our objective was to characterize the relationship between epicormics and provenance on 13-year-old Quercus petraea trees, taking into account their growth traits.

? On the growth units (GU) studied, epicormics were essentially isolated buds (95%) and they were mainly located at branch bases, secondarily on GU limits and marginally on lateral position. Provenance effect was very small on the number of epicormics and slightly larger on the number of branches and GU limits. On the contrary, the growth situation-independently from provenance-proved to have a larger effect on the 2 latter growth traits.

? Altogether, these results clearly show the predominant effect of morphological traits and thus of ontogeny. This opens important research perspectives: how growth conditions and genetics (at progeny and individual levels) influence height growth, setting and fate of axillary buds on the one hand and fate of epicormic buds inserted directly on the stem, or at branch bases on the other hand.

     

Properties of boundary-line release criteria in North American tree species

? Boundary line release criteria are increasingly applied to evaluate forest disturbance histories from tree-ring data. However, a number of important properties central to the technique have not been evaluated, including: (i) the ability of boundary line release criteria to standardize releases across various sites, species, and tree life stages (ii) the minimum sample sizes necessary for developing boundary lines, and (iii) the degree to which the criteria can resolve the degree of crown exposure following a disturbance event.

? In an analysis of eleven North American tree species, boundary line release criteria do not fully compensate for declines in release response a tree experiences with increasing age and size, with the exception Tsuga canadensis.

? A bootstrapping analysis indicates that approximately 50 000 ring width measurements are necessary to develop boundary line release criteria for a given species.

? In a Quercus prinus stand, boundary line release criteria better predict the degree of crown exposure following a disturbance than an earlier running mean technique.

? Despite certain limitations, boundary line release criteria have the potential to standardize release calculation across most life stages of a tree, and possibly among sites and species.