Photosynthetic responses to CO2 enrichment of four hardwood species in a forest understory

We compared the CO₂- and light-dependence of photosynthesis of four tree species (Acer rubrum, Carya glabra, Cercis canadensis, Liquidambar styraciflua) growing in the understory of a loblolly pine plantation under ambient or ambient plus 200 μl l^–1 CO₂. Naturally-established saplings were fumigated with a free-air CO₂ enrichment system. Light-saturated photosynthetic rates were 159–190% greater for Ce. canadensis saplings grown and measured under elevated CO₂. This species had the greatest CO₂ stimulation of photosynthesis. Photosynthetic rates were only 59% greater for A. rubrum saplings under CO₂ enrichment and Ca. glabra and L. styraciflua had intermediate responses. Elevated CO₂ stimulated light-saturated photosynthesis more than the apparent quantum yield. The maximum rate of carboxylation of ribulose-1,5-bisphosphate carboxylase, estimated from gas-exchange measurements, was not consistently affected by growth in elevated CO₂. However, the maximum electron transport rate estimated from gas- exchange measurements and from chlorophyll fluorescence, when averaged across species and dates, was approximately 10% higher for saplings in elevated CO₂. The proportionately greater stimulation of light-saturated photosynthesis than the apparent quantum yield and elevated rates of maximum electron transport suggests that saplings growing under elevated CO₂ make more efficient use of sunflecks. The stimulation of light-saturated photosynthesis by CO₂ did not appear to correlate with shade-tolerance ranking of the individual species. However, the species with the greatest enhancement of photosynthesis, Ce. canadensis and L. styraciflua, also invested the greatest proportion of soluble protein in Rubisco. Environmental and endogenous factors affecting N partitioning may partially explain interspecific variation in the photosynthetic response to elevated CO₂. Received: 16 February 1999 / Accepted: 30 August 1999     

Higher northern latitude normalized difference vegetation index and growing season trends from 1982 to 1999

Normalized difference vegetation index data from the polar-orbiting National Oceanic and Atmospheric Administration meteorological satellites from 1982 to 1999 show significant variations in photosynthetic activity and growing season length at latitudes above 35°N. Two distinct periods of increasing plant growth are apparent: 1982–1991 and 1992–1999, separated by a reduction from 1991 to 1992 associated with global cooling resulting from the volcanic eruption of Mt. Pinatubo in June 1991. The average May to September normalized difference vegetation index from 45°N to 75°N increased by 9% from 1982 to 1991, decreased by 5% from 1991 to 1992, and increased by 8% from 1992 to 1999. Variations in the normalized difference vegetation index were associated with variations in the start of the growing season of –5.6, +3.9, and –1.7 days respectively, for the three time periods. Our results support surface temperature increases within the same period at higher northern latitudes where temperature limits plant growth. Received: 25 October 2000 / Revised: 20 August 2001 / Accepted: 22 August 2001     

Levels of genetic diversity at different stages of the domestication cycle of interior spruce in British Columbia

Concerns over the reductionist nature of the domestication of forest-tree species focus on the possibility of potential genetic erosion during this process. To address these concerns, genetic diversity assessments in a breeding zone the Province of British Columbia “interior” spruce (Picea glauca×engelmanni) program was conducted using allozyme markers. Genetic-variation comparisons were made between natural and production (seed orchard) populations as well as seed and seedling crops produced from the same breeding zone’s seed orchard. The natural population sample consisted of a total of 360 trees representing three stands within each of three watersheds present in the Shuswap-Adams low-elevation zone of interior British Columbia. Small amounts of genetic differentiation were observed among the nine natural populations (4%) and this was attributable to extensive gene flow Consequently, the sum of these nine populations was considered as a baseline for the genetic variation present in the breeding zone. The comparisons between the seed orchard and the breeding zone produced a similar percentage of polymorphic loci while the expected hetrozygosity (0.207 vs 0.210) and the average number of alleles per locus (2.7 vs 2.4) were slightly lower in the seed orchard. A total of seven natural populations’ rare alleles were not present in the orchard population, while one allele was unique to the orchard. The %P increased to 70.6% in the seedlot, but dropped to the natural populations level (64.7%) in the plantation. The observed increase in %P was a result of pollen contamination in the orchard. It is suspected that the reduction in the plantation was caused by an unintentional selection in the nursery. Simulated roguing in the orchard did not drastically reduce even if up to 50% of the orchard’s clones were rogued. However, roguing was associated with a reduction in the average number of alleles per locus (i.e., sampling effect). Received: 2 January 1996 / Accepted: 24 May 1996     

Geographic pattern of genetic variation in the fox tapeworm Echinococcus multilocularis

Intraspecific genetic variation of Echinococcus multilocularis, the etiologic agent of human alveolar echinococcosis, has been evaluated among 76 geographic isolates from Europe, Asia and North America by using sequence data of mitochondrial and nuclear DNA. Relatively low genetic variation was found only in the mitochondrial DNA sequence consisting of 3 protein-coding genes. Pairwise divergence among the resultant 18 haplotypes ranged from 0.03 to 1.91%. Phylogenetic trees and parsimony network of these haplotypes depicted a geographic division into European, Asian and North American clades, but 1 haplotype from Inner Mongolia was unrelated to other haplotypes. The coexistence of the Asian and North American haplotypes could be seen, particularly on the St. Lawrence Island in the Bering Sea. These data suggest an evolutionary scenario in which distinct parasite populations derived from glacial refugia have been maintained by indigenous host mammals. The nuclear DNA sequence for the immunodominant B cell epitope region of ezrin/radixin/moesin-like protein (elp) was extremely conservative, indicating that the elp antigen is available for immunodiagnosis in any endemic areas.     

Effects of seasonal rainfall on radial growth in two tropical tree species

 Seasonal drought may limit primary productivity in most of the tropics, but the determinants of tree growth are not well known. A 10-year study of the deciduous trees Cochlospermum vitifolium (Willd.) Spreng. (Cochlospermaceae) and Cnidoscolus spinosus Lundell (Euphorbiaceae) in southwestern México showed radial growth to be highly correlated (both r>0.85) only with precipitation during an interval of <2 months in the mid-wet season. Growth was not affected by total annual precipitation or by an early starting or late ending of the wet season, or by heavy rainfall in the dry season. Annual mean girth increments ranged from 0.03 to 3.31 cm and −0.1 to 2.01 cm, respectively. The best model for growth (r ²>0.85) was a linear combination of mid-summer precipitation (positive coefficient) and total precipitation over the previous 2 years (negative coefficient). Comparison with other species showed heterogeneous responses of wood production to climate variation, and suggests that the range of functional types of dry forest trees is still unknown. Received: 9 September 1996 / Accepted: 4 March 1997     

Inter- and intrapopulation genetic variation in species ofPorphyra (Rhodophyta: Bangiales) from British Columbia and adjacent waters

Starch gel electrophoresis of 17 proteins has provided data on inter- and intrapopulation genetic variation in 20 species ofPorphyra occurring in British Columbia and adjacent areas.P. cuneiformis andP. nereocystis showed no within species variation, even over ranges of more than 1000 km. Populations ofP. abbottae, P. fallax, P. fucicola, P. gardneri andP. schizophylla were characterized by fixation for certain alleles. The number of polymorphic loci in a population ranged from zero to nine, depending on the species. Six species had populations that were polymorphic at just a single locus. Only two species (P. mumfordii andP. pseudolanceolata) had populations that were polymorphic at more than three loci. These levels of genetic variation are lower than those reported for populations of JapanesePorphyra species. Eleven taxa were polymorphic for 6-phosphogluconate dehydrogenase, the most variable enzyme. No within species polymorphisms were detected for bromoperoxidase, lactate dehydrogenase, superoxide dismutase or phycoerythrin. Possible evidence for the chimeric nature of the thallus was observed only inP. mumfordii.     

Geographic variation and dispersal history in Fennoscandian populations of two forest herbs

 Carex digitata and Melica nutans are forest understorey herbs with wide European distributions and their northern range margins in Fennoscandia. The species have closely similar habitat requirements, occur in small populations in old forest stands on base-rich to neutral soils and have restricted dispersal abilities at the present day. This study investigates the structure of allozyme variation (12 and 8 loci, respectively) in material of both species (38 and 37 populations, respectively) from throughout southern Sweden and southern Finland. Both species show a relatively low overall genetic diversity (H_T excluding monomorphic loci=0.17 and 0.18, respectively). The hierarchic structuring of allelic diversity in the species is similar, with a relatively high between-population component of diversity (G_ST=0.36 and 0.37, respectively). Neither of the species shows a clear intraspecific pattern of geographic differentiation. The lack of large-scale patterns of geographic differentiation is not consistent with a simple scenario of discrete and independent waves of immigration into Fennoscandia. However, particularly in M. nutans, a group of populations from a lowland belt across southwestern Finland and southern central Sweden is somewhat differentiated from populations to the north and south. A number of rare alleles in both species are widely, but patchily distributed in low frequencies. Hybridization may account for the scattered occurence of some of the rare alleles in Carex digitata, but cannot explain the distribution of rare alleles in Melica nutans. Received July 23, 2001 Accepted December 6, 2001     

Characteristics of genetic variation in the progenies of protoplast-derived plants of rice, Oryza sativa cv Nipponbare

Genetic variation in protoplast-derived rice (Oryza sativa L.) plants was characterized using first and second generation selfed progenies. A total of 133 regenerated plants were obtained from ten protoplasts of the japonica rice cultivar Nipponbare. Sixty two regenerated plants which set enough seeds for the subsequent field tests at the next generation and were derived from five protoplasts were selected, and their selfed seeds were used as the first selfed-seed progeny generation). Fifteen plants were selected from each of the 15 lines, and their selfed seeds were used for tests at the generation. Thirty seven lines (60%) segregated plants with detrimental mutant characters of yellow-green phenotype, dwarf stature, dense and short panicle, or low seed fertility. According to the segregation patterns in the lines having mutated plants among those originated from the same protoplasts, the stages of mutation induction were estimated. Additionally, five quantitative traits were changed in almost all and lines. Varied quantitative traits of heading date, number of spikelets per panicle, and seed fertility, were in a heterozygous state. However, culm and panicle lengths showed high uniformity, whereas reduced culm and panicle lengths were caused by mutational changes in polygenes and/or multiple genes. Received: 20 March 1996 / Accepted: 21 June 1996     

Reproductive system and pattern of genetic variation in two Limnanthes species

Summary Several populations of two species of the genus Limnanthes, (L. alba, an outbreeder and L. floccosa, an inbreeder) were examined with respect to variability of fifteen quantitative characters, allozyme variation at 11 loci, and response to different pollination conditions and moisture stress. Nearly equal amounts of phenotypic variability were found in the two species. L. alba had higher within-family variability than L. floccosa, but this result was highly heterogeneous among characters. A study of between- and within-population variance estimates did not reject the null hypothesis that L. alba and L. floccosa are similar with regard to the partitioning and amount of variability for quantitative characters. However, allozyme variation at 11 loci in a large number of populations showed L. alba to be highly polymorphic in contrast to the virtual monomorphism within L. floccosa populations. The average number of alleles per locus in L. alba and L. floccosa was 1.97 and 1.02, respectively, and on an average, L. alba and L. floccosa populations had 63% and 3% loci with polymorphism, respectively. Three groups of allozyme allelic combinations emerged which correlated well with the taxonomic delineation of allogamous L. alba, three semi-autogamous L. floccosa forms and two autogamous L. floccosa forms.All taxa showed a significant reduction in the seed output per plant due to moisture stress. L. alba suffered a further loss of fecundity under the paucity of pollinators, L. floccosa ssp. floccosa showed no significant effect from this factor, whereas L. floccosa ssp. grandiflora exhibited a curvilinear response which peaked at partial pollination and decreased to a lower level at full pollination.The geographic distribution of the two species with regard to the temperature and rainfall distribution did not suggest L. floccosa to be living in drier marginal areas. Patterns of variation in flowering time showed L. alba to be less variable than L. floccosa. Overall, there seemed to be little direct support for the thesis that inbreeding species originated from outcrossing taxa in marginal environments as a direct adaptation to a shortened growing season of xeric environments and to the lack of pollinators. Alternative hypotheses suggest that autogamy in L. floccosa might have evolved as a reproductive isolating barrier acting through either cleistogamy or divergence in flowering times.     

The importance of phenology for the evaluation of impact of climate change on growth of boreal, temperate and Mediterranean forests ecosystems: an overview

An overview is presented of the phenological models relevant for boreal coniferous, temperate-zone deciduous and Mediterranean coniferous forest ecosystems. The phenology of the boreal forests is mainly driven by temperature, affecting the timing of the start of the growing season and thereby its duration, and the level of frost hardiness and thereby the reduction of foliage area and photosynthetic capacity by severe frost events. The phenology of temperate-zone forests is also mainly driven by temperature. Since temperate-zone forests are mostly mixed-species deciduous forests, differences in phenological response may affect competition between tree species. The phenology of Mediterranean coniferous forests is mainly driven by water availability, affecting the development of leaf area, rather than the timing of phenological events. These phenological models were subsequently coupled to the process-based forest model FORGRO to evaluate the effect of different climate change scenarios on growth. The results indicate that the phenology of each of the forest types significantly affects the growth response to a given climate change scenario. The absolute responses presented in this study should, however, be used with caution as there are still uncertainties in the phenological models, the growth models, the parameter values obtained and the climate change scenarios used. Future research should attempt to reduce these uncertainties. It is recommended that phenological models that describe the mechanisms by which seasonality in climatic drivers affects the phenological aspects of trees should be developed and carefully tested. Only by using such models may we make an assessment of the impact of climate change on the functioning and productivity of different forest ecosystems. Received: 21 October 1999 / Revised: 10 May 2000 / Accepted: 10 May 2000     

Genetic variation in low elevation Douglas-fir of British Columbia and its relevance to gene conservation

Patterns of variation were studied at 20 isozyme loci in 49 coastal, low-elevation Douglas-fir populations in SW British Columbia and NW Washington State. Several components of variation were estimated for each population including the number of alleles per locus N _a, number of alleles per polymorphic locus N _a(95), inbreeding F, heterozygosity H, and population divergence D. F was near zero indicating nearly complete outcrossing within populations. H was quite high (16%) and in aecord with previous studies of Douglas-fir. D values were low (equivalent to Wrights F _ST of 0.08) indicating levels of gene flow sufficient to largely homogenize populations. The parameters of diversity N _a, N _a(95), H, and D showed little intercorrelation across populations. A homogenous pattern of genetic relationship among populations was shown by the clustering of populations based on their inferred relationship, and by the principal components of the matrix of inferred genetic relationship. Because of the complex nature of gene diversity and the continuous nature of population differentiation in Douglas-fir, it is difficult with isozyme markers to identify specific populations of value for genetic conservation in this species.This paper is dedicated to J.C. Heaman on the occasion of his retirement.     

Geographic distribution and multilocus organization of isozyme variation of rice (Oryza sativa L.)

Genetic organization of isozyme variation in rice (Oryza sativa L.) was investigated based on 17 polymorphic isozyme loci using a sample of 511 accessions of worldwide origin. The genetic diversity within the species was very high (H=0.36 with 4.82 alleles per locus), as compared with most selfing plant species. Three diversity centers were detected for isozyme variation including South Asia, China and Southeast Asia. The accessions were classified into three well-differentiated cultivar groups corresponding to the indica and japonica subspecies, and a new unnamed group. Variation within the cultivar groups accounted for 80% of the total isozyme variation. Within-country variation accounted for 58% of the total variation while among-region and among-country variation within the cultivar groups accounted for only 14% and 8% of the total variation. Analyses using log-linear models revealed that pronounced non-random associations between and among alleles at many unlinked isozyme loci were organized in a non-hierarchical pattern, and subspecific and macro-geographic differentiation was much more pronounced in multilocus phenotype frequencies than in allelic frequencies at individual loci. These results suggest that selection on multilocus gene complexes was largely responsible for the maintenance of the extensive isozyme variation within the species and the indica-japonica differentiation. Our results further suggest the independent domestication of indica and japonica, the dual origins of the indica rice from China and South Asia (India), and the differentiation of the ecotypes ’javanica’ and the ’temperate japonica’ within the japonica subspecies. Received: 5 August 1999 / Accepted: 13 December 1999     

Phenological stages of natural species and their use as climate indicators

 The objectives of this paper are to: (1) present 10 years of phenological data for nine natural species growing in a Mediterranean-type climate, (2) present threshold temperatures that were derived for the computation of cumulative degree-days (CDD), and (3) evaluate the sensitivity of the nine natural species to weather variability. The study was conducted at the Phenological Research Garden of Oristano, Sardinia, Italy, during the period 1986–96. The observations were made on five typical Mediterranean species and four species that are typical of higher latitudes. The mean annual pattern of phenological events and the CDD from 1 January are given for each development stage. Temperature thresholds were evaluated by comparing the standard deviation about the mean number of days in the development period for each species. A good relationship between timing of phenophase occurrence and temperature was observed for the Mediterranean species, which were little affected by variations in rainfall. Phenological development of the non-native species was affected by springtime rainfall. Accepted: 28 October 1998     

Trends in phenological phases in Europe between 1951 and 1996

Increases in air temperature due to the anthropogenic greenhouse effect can be detected easily in the phenological data of Europe within the last four decades because spring phenological events are particularly sensitive to temperature. Our new analysis of observational data from the International Phenological Gardens in Europe for the 1959–1996 period revealed that spring events, such as leaf unfolding, have advanced on average by 6.3 days (–0.21 day/year), whereas autumn events, such as leaf colouring, have been delayed on average by 4.5 days (+0.15 day/year). Thus, the average annual growing season has lengthened on average by 10.8 days since the early 1960s. For autumn events, differences between mean trends of species could not be detected, but for spring events there were differences between species, with the higher trends for leaf unfolding and flowering of shrubs indicating that changes in events occurring in the early spring are more distinct. These observed trends in plant phenological events in the International Phenological Gardens and results of other phenological studies in Europe, summarised in this study, are consistent with AVHRR satellite measurements of the normalized difference vegetation index from 1981 to 1991 and with an analysis of long-term measurements of the annual cycle of CO₂ concentration in Hawaii and Alaska, also indicating a global lengthening of the growing season. Received: 21 October 1999 / Accepted: 2 March 2000     

Evaluation of genetic variation in the daylily (Hemerocallis spp.) using AFLP markers

The daylily (Hemerocallis spp.) is one of the most economically important ornamental plant species in commerce. Interestingly, it is also one of the most heavily bred crops during the past 60 years. Since the American Hemerocallis Society began acting as the official registry of daylily cultivars in 1947, more than 40 000 registrations have been processed. In order to determine the effects of intensive breeding on cultivar development, and to study relationships among different species, genetic variation in the daylily was estimated using AFLP markers. Nineteen primary genotypes (species and early cultivars) and 100 modern cultivars from different time periods were evaluated using 152 unambiguous bands (average 79% polymorphism rate) derived from three AFLP primer combinations. Overall, pairwise similarity estimates between entries ranged between 0.618 and 0.926 (average=0.800). When comparing cultivar groups from different time periods (1940–1998), genetic similarity was initially increased, compared to the primary diploid genotypes, remained constant from 1940 to 1980, and then steadily increased as breeding efforts intensified and hybridizers began focusing on a limited tetraploid germplasm pool derived by colchicine conversion. Among modern (1991–1998) daylily cultivars, genetic similarity has increased by approximately 10% compared to the primary genotypes. These data were also used to evaluate recent taxonomic classifications among daylily species which, with a few minor exceptions, were generally supported by the AFLP data. Received: 15 March 2000 / Accepted: 13 June 2000     

Allozyme variation in Carex sect. Digitatae – Evidence of introgression, genetic distinctiveness and evolution of taxa

 A subset of populations of the European taxa of Carex sect. Digitatae, including Carex digitata, C. pallens, C. ornithopoda, C. pediformis ssp. rhizodes and C. humilis have been investigated for allozyme variation. The (presumably) distantly related C. pendula was used as a reference taxon. Carex digitata, C. pallens and C. ornithopoda on the one hand, and C. humilis and C. pediformis on the other, shared the majority of the alleles. Cluster analyses based on genetic distances grouped populations of C. digitata and C. ornithopoda in a mixed subcluster whereas all populations of C. pallens formed a subcluster distinct from the digitata/ornithopoda cluster. This in spite of the fact that C. ornithopoda is morphologically clearly distinct from C. digitata whereas C. pallens is barely distinguishable from the latter. Carex pediformis and C. humilis appeared not more genetically similar to C. digitata than to the reference taxon. It is concluded that 1) C. digitata, C. pallens and C. ornithopoda are closely related and most probably forms a monophyletic group, 2) C. pallens is a genetically distinct species, 3) C. pediformis and C. humilis are not closely related to the rest of C. sect. Digitatae, 4) C. pediformis and C. humilis are relatively closely related, 5) introgression do occur between the investigated species but not to such an extent that it can explain major geographic patterns of variation in C. digitata. Received December 21, 2001; accepted November 14, 2002 Published online: March 24, 2003     

Application of a progressive-difference method to identify climatic factors causing variation in the rice yield in the Yangtze Delta, China

Time series of rice yields consist of a technology-driven trend and variations caused by climate fluctuations. To explore the relationship between yields and climate, the trend and temporal variation often have to be separated. In this study, a progressive-difference method was applied to eliminate the trend in time series. By differentiating yields and climatic factors in 2 successive years, the relationship between variations in yield and climatic factors was determined with multiple- regression analysis. The number of hours of sunshine, the temperature and the precipitation were each defined for different intervals during the growing season and used as different regression variables. Rice yields and climate data for the Yangtze Delta of China from 1961 to 1990 were used as a case study. The number of hours of sunshine during the tillering stage and the heading to milk stage particularly affected the yield. In both periods radiation was low. In the first period, the vegetative organs of the rice crop were formed while in the second period solar radiation was important for grain filling. The average temperature during the tillering to jointing stage reached its maximum, which affected rice yields negatively. Precipitation was generally low during the jointing and booting stages, which had a positive correlation with yield, while high precipitation had a negative effect during the milk stage. The results indicate that the climatic factors should be expressed as 20- to 30-day averages in the Yangtze Delta; a shorter or longer period, e.g. 10 or 40 days, is less appropriate. Received: 30 May 2000 / Revised: 27 October 2000 / Accepted: 30 October 2000     

Allozyme variation in Mexican species and classification of Datura (Solanaceae)

 Fifty natural Datura populations, belonging to eleven species (D. ceratocaula, D. discolor, D. inoxia, D. kymatocarpa, D. lanosa, D. metel, D. pruinosa, D. quercifolia, D. reburra, D. stramonium, D. wrightii) from Mexico and adjacent USA, were investigated using starch gel electrophoresis. A total of 64 alleles were scored at 17 loci (DIA1, DIA2, GOT1, GOT2, G6PDH, IDH, MDH1, MDH2, MDH3, ME, PGD1, PGD2, PGM1, PGM2, PHI, SAD, SOD). The heterozygosity among the species ranged from 0.166 (D. ceratocaula) to 0.276 (D. wrightii). Most genetic diversity was found within populations (average H_s=0.242), while values between populations are relatively low (average Dst=0.066, Gst=0.171). The analysis of the genetic distance suggested new taxonomic relationships among the species. Rather than supporting the conventional infrageneric classification with three sections, the results revealed that the herbaceous members of the genus Datura form four groups. One group included four of the eight species of the section Dutra and was more similar to the section Ceratocaulis than it was to the other group that contained the remaining taxa of Dutra. Received February 13, 2001 Accepted December 25, 2001     

Determining the growing season of land vegetation on the basis of plant phenology and satellite data in Northern China

The objectives of this study are to explore the relationships between plant phenology and satellite-sensor-derived measures of greenness, and to advance a new procedure for determining the growing season of land vegetation at the regional scale. Three phenological stations were selected as sample sites to represent different climatic zones and vegetation types in northern China. The mixed data set consists of occurrence dates of all observed phenophases for 50–70 kinds of trees and shrubs from 1983 to 1988. Using these data, we calculated the cumulative frequency of phenophases in every 5-day period (pentad) throughout each year, and also drew the cumulative frequency distribution curve for all station-years, in order to reveal the typical seasonal characteristics of these plant communities. The growing season was set as the time interval between 5% and 95% of the phenological cumulative frequency. Average lengths of the growing season varied between 188 days in the northern, to 259 days in the southern part of the research region. The beginning and end dates of the surface growing season were then applied each year as time thresholds, to determine the corresponding 10-day peak greenness values from normalized difference vegetation index curves for 8-km² pixels overlying the phenological stations. Our results show that, at the beginning of the growing season, the largest average greenness value occurs in the southern part, then in the northern, and finally the middle part of the research region. In contrast, at the end of the growing season, the largest average greenness value is measured in the northern part, next in the middle and lastly the southern part of the research region. In future studies, these derived NDVI thresholds can be applied to determine the growing season of similar plant communities at other sites, which lack surface phenological data. Received: 29 November 1999 / Revised: 14 March 2000 / Accepted: 15 March 2000     

Responses of canopy duration to temperature changes in four temperate tree species: relative contributions of spring and autumn leaf phenology

While changes in spring phenological events due to global warming have been widely documented, changes in autumn phenology, and therefore in growing season length, are less studied and poorly understood. However, it may be helpful to assess the potential lengthening of the growing season under climate warming in order to determine its further impact on forest productivity and C balance. The present study aimed to: (1) characterise the sensitivity of leaf phenological events to temperature, and (2) quantify the relative contributions of leaf unfolding and senescence to the extension of canopy duration with increasing temperature, in four deciduous tree species (Acer pseudoplatanus, Fagus sylvatica, Fraxinus excelsior and Quercus petraea). For 3 consecutive years, we monitored the spring and autumn phenology of 41 populations at elevations ranging from 100 to 1,600 m. Overall, we found significant altitudinal trends in leaf phenology and species-specific differences in temperature sensitivity. With increasing temperature, we recorded an advance in flushing from 1.9 ± 0.3 to 6.6 ± 0.4 days °C⁻¹ (mean ± SD) and a 0 to 5.6 ± 0.6 days °C⁻¹ delay in leaf senescence. Together both changes resulted in a 6.9 ± 1.0 to 13.0 ± 0.7 days °C⁻¹ lengthening of canopy duration depending on species. For three of the four studied species, advances in flushing were the main factor responsible for lengthening canopy duration with increasing temperature, leading to a potentially larger gain in solar radiation than delays in leaf senescence. In contrast, for beech, we found a higher sensitivity to temperature in leaf senescence than in flushing, resulting in an equivalent contribution in solar radiation gain. These results suggest that climate warming will alter the C uptake period and forest productivity by lengthening canopy duration. Moreover, the between-species differences in phenological responses to temperature evidenced here could affect biotic interactions under climate warming.