Relative growth rate of leaf biomass and leaf nitrogen content in several mediterranean woody species

In many plant species, herbivory is a major determinant of leaf mortality and it can cause a strong reduction in productive potential. Most predation occurs on young, expanding leaves. Thus, a rapid growth of the leaves can reduce the impact of predation. Furthermore, in cold Mediterranean climates the length of the growing season is constrained to a short period in spring and early summer owing both to low winter temperatures and drought stress in early summer. Therefore, a rapid deployment of leaf area and a high photosynthetic capacity during the spring and early summer might have important positive effects on the final carbon balance of the leaf population. Relative growth rates (RGR) of leaf biomass were measured in 19 woody species typical of Central Western Spain with deciduous and evergreen habits. Highly significant differences were detected in the leaf growth rate of the different species. The differences between species, however, did not correlate either with the mean leaf life-span of each of the species or with other leaf traits such as photosynthetic capacity, specific leaf area or nitrogen content. Leaf growth rate was positively correlated with time elapsed between leaf initiation and fruit maturation, so that species with fruit dispersal in spring and early summer in general had lower leaf growth rates than species with autumn fruit shedding. This relationship shows the effects of the concurrence between vegetative and reproductive organs for nutrients and other resources. Nitrogen concentration in the leaves was very high at the time of bud break, and declined during leaf expansion owing to the dilution associated with the increase in structural components. The rate of nitrogen dilution was, thus, positively related to the leaf growth rate. Relative growth rates calculated for nitrogen mass in leaves were very low compared to the growth in total mass. This suggests that most leaf nitrogen is translocated from the plant stores to the leaf biomass before the start of leaf expansion and that the contribution of root uptake during leaf expansion is comparatively low.     

Seasonal Changes in the Physiology of S24 Perennial Ryegrass (Lolium perenne L.). 2. Potential Leaf and Canopy Photosynthesis during the Transition from Vegetative to Reproductive growth

The photosynthetic potential of successive youngest fully-expandedleaves of S24 L. perenne, grown as simulated swards under naturalenvironmental conditions, was measured during establishmentin autumn, over winter and during the transition from vegetativeto reproductive growth the following spring. Measurements weremade at a standard light energy receipt of 250 J m^–2 s^–1(400–700 nm) and at 15 °C. The photosynthetic potential of the leaves decreased in autumnas the swards increased in density under worsening environmentalconditions. During the spring, photosynthetic rates rose fromlow over-winter values so that by March, before stem elongationbegan, they were equal to the rates in the previous autumn.Following stem elongation there was a further increase in leafpotential. Reasons for these changes in leaf potential are discussed. During spring, the photosynthetic potential of the canopy alsorose - both as measured, and as predicted by the Monteith modelof canopy photosynthesis. Use of the model suggested that increasingleaf potential made the greatest contribution to the rise inthe potential of the canopy, although, following stem elongation,changes in LAI and canopy structure had a further significanteffect.     

Sources ofCycloconium oleaginum (Cast.) conidia for infection of olive leaves and conditions determining leaf spot disease development in the region of Sétif,Algeria

In the region of Sétif, peacock leaf spot disease caused byCycloconium oleaginum was found to be most prevalent in the period from late autumn to spring and of minor significance in the period from the beginning of July until the middle of November. Severity of infection on the lower parts was greater than on the upper parts of the trees. Damage on leaves facing north was much greater than on those facing south. Production of conidia leaf spots was found to be high in spring and late autumn but very low in summer and early autumn. Temperatures from 15 to 18°C were found optimal for the growth of the fungus. Reduced growth was seen at 3, 1 and 25°C with total inhibition at 30°C. Our results suggest that fallen leaves play no role in new infections and the role of the remaining spots on the tree during summer is of little importance. Four phases for the infection of new leaves were determined. In the first, during late spring, three newly opened pairs of leaves were infected, this infection remains hidden until late autumn. The second phase occurs in early autumn after rain. The third stage in late autumn and in the beginning of winter is characterized by the occurrence of new leaf spots which are usually concentrated on the basal pair of newly grown leaves. The fourth phase of infection, at the beginning of spring, is the most important of all. The infected leaves at this stage, comprise the infection source for all the following stages.     

Morphological and phenological characteristics of leaf development ofDurio zibethinus Murray (Bombacaceae)

The morphological and phenological characteristics of leaf development ofDurio zibethinus Murray were investigated at an experimental field of Universiti Pertanian Malaysia (UPM) in Selangor. Proportionality was observed in the relations of leaf length to leaf width and of leaf area to the product of leaf width and length. The proportionality was explained from the similarity of leaf shape. New leaves emerged continuously, but the number of new leaves fluctuated seasonally. The emergence of leaves was inhibited by the flower bud formation. In the survival curves of leaves, the relative fall rate was lower at the early stage of leaf development than at the late stage. Leaf longevity of 100 to 133 days was low and leaf expansion period of two weeks was short in comparison with the published data on tropical trees. From the ecophysiological viewpoint, the leaf survival strategy of the present species was discussed: the present species manages to set up a photosynthetic system in a short period by the rapid leaf growth; the lower leaf longevity is advantageous to reaching more frequently high photosynthetic production by newly emerged leaves.     

Matter-economical roles of evergreen leaves inAucuba japonica,an understory shrub in the warm-temperate region of Japan

Seasonal dynamics in nitrogen and phosphorus content were examined for each component organ ofAucuba japonica, an evergreen understory shrub in the warmtemperate region of Japan. Evergreen foliage was the largest pool for each nutrient; nitrogen and phosphorus were accumulated and stored in autumn and then redistributed in the spring. For individual leaves, such seasonal accumulations and redistributions were repeated through two or three years and then at leaf fall, an additional amount was withdrawn. Rapid growth of new shoots and flowers during spring was supported by the massive redistribution of the nutrients from the old foliage. The redistribution accounted for 85% and 65% of the total nitrogen and phosphorus input to the new shoots, respectively. Such a high ratio of redistribution resulted in a conservative nutrient economy, and must be positively related to the photosynthetic production in the ligh-limited environment.     

Seasonal branch nutrient dynamics in two Mediterranean woody shrubs with contrasted phenology

BACKGROUND AND AIMS: Mediterranean woody plants have a wide variety of phenological strategies. Some authors have classified the Mediterranean phanaerophytes into two broad phenological categories: phenophase-overlappers (that overlap resource-demanding activities in a short period of the year) and phenophase-sequencers (that protract resource-demanding activities throughout the year). In this work the impact of both phenological strategies on leaf nutrient accumulation and retranslocation dynamics at the level of leaves and branches was evaluated. Phenophase-overlappers were expected to accumulate nutrients in leaves throughout most of the year and withdraw them efficiently in a short period. Phenophase-sequencers were expected to withdraw nutrients progressively throughout the year, without long accumulation periods. METHODS: To test this hypothesis, variations in phenology and leaf NPK in the crown of a phenophase-overlapper Cistus laurifolius and a phenophase-sequencer Bupleurum fruticosum were monitored monthly during 2 years. KEY RESULTS: Changes in nutrient concentration at the leaf level were not clearly related with the different phenologies. Nitrogen and phosphorous resorption efficiencies were lower in the phenophase-overlapper, and accumulation-retranslocation seasonality was similar in both species. Changes in the branch nutrient pool agreed with the hypothesis that the phenophase-overlapper accumulated nutrients from summer until the bud burst of the following spring, recovering a large nutrient pool during massive leaf shedding. The phenophase-sequencer did not accumulate nutrients from autumn until early spring, achieving lower nutrient recovery during spring leaf shedding. CONCLUSIONS: It is concluded that phenological demands influence branch nutrient cycling. This effect is easier to detect by assessing changes in the branch nutrient pool rather than changes in the leaf nutrient concentration.     

Seasonal change in specific needle weight ofPinus thunbergii

To evaluate the reserve dynamics in pine needles, an index of specific needle weight (SNW; dry weight per unit length of needles) was provided. Developed needles ofPinus thunbergii did not show any additional elongation in their second year. Thus the seasonal change in SNW of developed needles was proved to indicate the dynamics of reserves in needles. SNW showed distinct seasonal change. It increased from autumn to carly spring, followed by a decrease by mid summer. This pattern of change was similar to the change in specific leaf weight (SLW) of evergreen broad leaves in temperate regions. It was concluded that the change in SNW indicated clearly the dynamics of reserves in needles, showing the accumulation in non-growth seasons and the consumption in subsequent growth seasons. The decrease in SNW during the growth season was 10–15% of the maximum found in early spring.     

Distribution and leaf growth of Thalassodendron pachyrhizum den Hartog in Southern Western Australia

The seagrass Thalassodendron pachyrhizum den Hartog grows on limestone reef platforms. Monthly leaf biomass was measured over 2 years and showed a strong seasonal variation with maximum biomass of 500 g m⁻². This seagrass loses all its leaves except for a bud and this characteristic was used to obtain a conservative estimate of productivity by change in standing stock. Leaf growth during the growing season was 6.6 mg Cg⁻¹ day⁻¹. Leaf length frequencies showed that new leaves formed during autumn (March–April). They grew from autumn until spring (November) and began to senesce in summer, followed by leaf fall in late summer (February–March).The growth of rhizome shoots “invading” free substratum space and the growth of new stems was measured for a 300-day period; about 9 leaves were produced in this period.     

Matter-economical roles of the evergreen foliage ofAucuba japonica,an understory shrub in the warm-temperate region of Japan

Leaf demography and productivity ofAucuba japonica, an understory shrub in the warm-temperate region, were examined and dry matter economy was analyzed to evaluate the roles of the evergreen foliage. Turnover of leaves occurred during a short period in spring. The mean leaf life span was about 2.6 years. Annual NAR (net assimilation rate) of each sample shoot was calculated from the biomass and the total dead mass estimated from scars of leaves and floral parts. The average NAR was 1.34±0.22 g·g⁻¹·yr⁻¹. The ratio of dry matter produced by leaves during their whole life span to the initial investment was 3.45±0.37. The annual NAR calculated for individual plants was negatively related to the life span of their leaves. The seasonal change in SLW (specific leaf weight) showed that the reserve material in leaves was accumulated from autumn to early spring and was consumed for the growth of new organs in the following season. The dry matter withdrawn in spring from the overwintering foliage amounted to 40% of dry mass of the new organs developed.     

Seasonal photosynthetic changes in the green-stemmed Mediterranean shrub <Emphasis Type="Italic">Calicotome villosa</Emphasis>: a comparison with leaves

Some photosynthetic attributes of leaves and stems were seasonally followed in the small-leaved, summer-deciduous, green-stemmed Mediterranean shrub Calicotome villosa. Both leaves and stems displayed similar photon energy-saturated photosystem 2 (PS2) efficiencies with a minimum during winter. A second minimum in stems during the leafless summer period could be ascribed to sustained photoinhibition. Yet, stems were slightly inferior in photon capture, resulting partly from lower chlorophyll (Chl) contents and partly from higher reflectance due to pubescence. As a result, photon energy-saturated linear electron transport rates were slightly higher in leaves. However, when the total leaf and stem areas were taken into account, this superiority was abolished during autumn and winter and more than overturned during spring. Given that during summer the stems were the only photosynthetic organs, the yearly photosynthetic contribution of stems was much higher. Chl contents in stems displayed a transient and considerable summer drop, accompanied by an increase in the carotenoid to Chl ratio, indicating a photo-protective adaptation to summer drought through a decrease of photo-selective capacity, typical for leaves of many Mediterranean plants.     

Grasstree (Xanthorrhoea preissii ) leaf growth in relation to season and water availability

Abstract Water stress usually arrests growth of even the most deep‐rooted species during summer drought in Mediterranean‐type climates. However, scant evidence suggests that grasstrees may represent an unusual exception. We used weather data and plant water potential to investigate the relationship between leaf growth and season in the grasstree, Xanthorrhoea preissii Endl. (Xanthorrhoeaceae). Leaf production in two contrasting habitats revealed continuous annual growth, oscillating between maximum rates (2.5–3.2 leaves/d) in late‐spring to autumn, to a minimum rate of 0.5 leaf/d during winter but never stopping. While the rate of leaf production during the fast‐growth season was positively correlated with temperature above 17–18°C, leaf elongation commenced substantially earlier in the year (from 12°C). Leaf water potentials cycled annually, with predawn readings commonly measured as zero during winter–spring and as low as ?1.26 MPa during summer, but never indicating stress by exceeding the turgor loss point. Leaf death was synchronized with summer drought. The fast (summer) growth period was characterized by rapidly fluctuating leaf production, particularly in banksia woodland, where plant growth reliably responded quickly to >18 mm of rainfall. Within 24 h of 59 mm of simulated rainfall, grasstrees in banksia woodland showed a marked increase in water potential, and leaf production reached 7.5 times the controls, confirming their capacity to respond to temporary spasmodic summer rains. Rainfall was the best climatic variable for predicting woodland grasstree leaf production during summer, whereas leaf production of forest grasstrees was most closely correlated with daylength. This plastic response of grasstrees between seasonal weather extremes is relatively rare among other mediterranean floras, and has implications for a recently proposed technique for ageing grasstrees.     

The leaf anatomy of a broad-leaved evergreen allows an increase in leaf nitrogen content in winter

In temperate regions, evergreen species are exposed to large seasonal changes in air temperature and irradiance. They change photosynthetic characteristics of leaves responding to such environmental changes. Recent studies have suggested that photosynthetic acclimation is strongly constrained by leaf anatomy such as leaf thickness, mesophyll and chloroplast surface facing the intercellular space, and the chloroplast volume. We studied how these parameters of leaf anatomy are related with photosynthetic seasonal acclimation. We evaluated differential effects of winter and summer irradiance on leaf anatomy and photosynthesis. Using a broad-leaved evergreen Aucuba japonica , we performed a transfer experiment in which irradiance regimes were changed at the beginning of autumn and of spring. We found that a vacant space on mesophyll surface in summer enabled chloroplast volume to increase in winter. The leaf nitrogen and Rubisco content were higher in winter than in summer. They were correlated significantly with chloroplast volume and with chloroplast surface area facing the intercellular space. Thus, summer leaves were thicker than needed to accommodate mesophyll surface chloroplasts at this time of year but this allowed for increases in mesophyll surface chloroplasts in the winter. It appears that summer leaf anatomical characteristics help facilitate photosynthetic acclimation to winter conditions. Photosynthetic capacity and photosynthetic nitrogen use efficiency were lower in winter than in summer but it appears that these reductions were partially compensated by higher Rubisco contents and mesophyll surface chloroplast area in winter foliage.     

Significance of sequential leaf development for nutrient balance of the cotton sedge,Eriophorum vaginatum L.

Summary The sedgeEriophorum vaginatum in an interior Alaskan muskeg site produced leaves sequentially at about 1.5-month intervals. Each leaf remained active for two growing seasons. Young leaves (even those initiated late in the season) always had high concentrations of N, P, K and Mg and were low in Ca. Stems had high concentrations of nutrients, sugar, amino acid N and soluble organic P in autumn and spring but low concentrations in summer. Growth of leaves in spring was strongly supported by translocation from storage. Leaves approached their maximum nutrient pool before nutrient uptake began in late spring, one month before maximum biomass. Retranslocation of nutrients from aging leaves could support nutrient input into new, actively growing leaves as a consequence of the sequential leaf development. For instance retranslocation from aging leaves accounted for more than 90 and 85% of P and N input to new leaves appearing in early summer and 100% to leaves that appeared later. Leaching losses were negligible. Half time for decay of standing dead litter was 10 years. We suggest that sequential leaf development paired with highly efficient remobilization of nutrients from senescing leaves enables plants to recycle nutrients within the shoot and minimize dependence upon soil nutrients. This may be an important mechanism enablingEriophorum vaginatum to dominate nutrient-poor sites. This may also explain why graminoids with sequential leaf production cooccur with evergreen shrubs and dominate over forbs and deciduous shrubs in nutrient-poor sites in the boreal forest (e.g., in bogs) and at the northern limit of the tundra zone.     

Differences in response to simulated herbivory between <Emphasis Type="Italic">Quercus crispula</Emphasis> and <Emphasis Type="Italic">Quercus dentata</Emphasis>

To evaluate the responses of Quercus crispula and Quercus dentata to herbivory, their leaves were subjected to simulated herbivory in early spring and examined for the subsequent changes in leaf traits and attacks by chewing herbivores in mid summer. In Quercus crispula, nitrogen content per area was higher in artificially damaged leaves than in control leaves. This species is assumed to increase the photosynthetic rate per area by increasing nitrogen content per area to compensate leaf area loss. In Quercus dentata, nitrogen content per area did not differ between artificially damaged and control leaves, while nitrogen content per mass was slightly lower in artificially damaged leaves. The difference in their responses can be attributable to the difference in the architecture of their leaves and/or the severeness of herbivory. The development of leaf area from early spring to mid summer was larger in artificially damaged leaves than in control leaves in both species, suggesting the compensatory response to leaf area loss. Leaf dry mass per unit area was also larger in artificially damaged leaves in both species, but the adaptive significance of this change is not clear. In spite of such changes in leaf traits, no difference was detected in the degree of damage by chewing herbivores between artificially damaged and controlled leaves in both species.     

Physiological and morphological acclimation of shade-grown tree seedlings to late-season canopy gap formation

Because acclimation to canopy gaps may involve coordination of new leaf production with morphological or physiological changes in existing, shade-developed leaves, we examined both new leaf production and photosynthesis of existing leaves on shade-grown seedlings after exposure to a late-season canopy gap. Midway through the summer, we transferred potted, shade-grown seedlings of four co-occurring temperate deciduous tree species representing a range of shade-tolerance categories and leaf production strategies to gaps. Shade-tolerant Acer saccharum was the least responsive to gap conditions. It produced few new, high-light acclimated leaves and increases in photosynthetic rates of shade-developed leaves appeared stomatally limited. Intermediately shade-tolerant Fraxinus americana and Quercus rubra responded most, by producing new leaves and increasing photosynthetic rates of existing shade-developed leaves to levels not significantly different from gap-grown controls within four weeks of gap exposure. Shade-intolerant Liriodendron tulipifera was intermediate in response. In these species, the degree of shoot-level morphological acclimation (new leaf production) and leaf-level physiological acclimation (photosynthetic increases in existing leaves) appear coupled. Mechanisms of acclimation also appear related to intrinsic patterns of nitrogen use and mobilization, the ability to adjust stomatal conductance, and shade tolerance.     

Carry-over effects of ozone and water stress on leaf phenological characteristics and bud frost hardiness of <Emphasis Type="Italic">Fagus crenata</Emphasis> seedlings

We examined the carry-over effects of ozone (O₃) and/or water stress on leaf phenological characteristics and bud frost hardiness of Fagus crenata seedlings. Three-year-old seedlings were exposed to charcoal-filtered air or 60 nl l^–1 O₃, 7 h a day, from May to October 1999 in naturally-lit growth chambers. Half of the seedlings in each gas treatment received 250 ml of water at 3-day intervals (well-watered treatment), while the rest received 175 ml of water at the same intervals (water-stressed treatment). All the seedlings were moved from the growth chambers to an experimental field on October 1999, and grown until April 2000 under field conditions. The exposure to O₃ during the growing season induced early leaf fall and reduction in leaf non-structural carbohydrates concentrations in the early autumn, as well as resulting in late bud break and reduction in the number of leaves per bud in the following spring. However, O₃ did not affect bud frost hardiness in the following winter. On the contrary, water stress did not affect leaf phenological characteristics, leaf and bud non-structural carbohydrates concentrations and bud frost hardiness. There were no significant synergistic or antagonistic effects of O₃ and water stress on leaf phenological characteristics, concentrations of leaf and bud non-structural carbohydrates and bud frost hardiness of the seedlings. These results show that the carry-over effects of O₃ can be found on the phenological characteristics and leaf non-structural carbohydrates concentrations, although there are almost no carry-over effects of water stress on phenological characteristics and winter hardiness of the seedlings.     

Growth Rings,Phenology, Hurricane Disturbance and Climate in Cyrilla racemiflora L., a Rain Forest Tree of the Luquillo Mountains,Puerto Rico1

The growth phenology of Cyrilla racemiflora L., the dominant tree species of the montane rain forest, (subtropical lower montane rain forest, sensu Holdridge) of the Luquillo Mountains of Puerto Rico was studied intensively during 1989, and then semiannually through mid-1993 to determine the periodicity of changes in xylem structure. Four trees at 770 m were monitored for flowering, branch elongation, leaf litterfall, and xylem cell growth and differentiation in the lower stem, and these events were related to local seasonal patterns of rainfall and temperature. Hurricane Hugo defoliated study trees in September, 1989. Bud-break and branch elongation in March, 1989 were followed by earlywood xylem cell production in the lower stem in April and the onset of flowering in May. Leaf litterfall was greatest between April and June, coinciding with peak branch growth and new leaf formation. Latewood xylem was produced in December. The general phenological pattern was synchronized between trees and over study years. Vessel diameter and density were monitored along with thickness of earlywood and latewood and the former converted to vessel lumen area, a measure of xylem conductance capacity. Annual growth rings were formed with periods of earlywood and latewood production coinciding with traditional summer (rainy) and winter (dry) seasons, respectively, in the Luquillo Mountains. Hurricane defoliation was followed by heavy flowering in 1990, a year of reduced branch elongation and annual xylem ring width, and was associated with maximum vessel lumen area, as was flowering in 1989, prior to the hurricane. Hurricane Hugo provided a perturbation that, through its elicited stress response, allowed for the demonstration of the interplay between flowering, branching, structural growth of xylem, and xylem function.     

Overwintering leaves of a forest-floor fern, Dryopteris crassirhizoma (Dryopteridaceae): a small contribution to the resource storage and photosynthetic carbon gain

BACKGROUND AND AIMS: Dryopteris crassirhizoma is a semi-evergreen fern growing on the floor of deciduous forests. The present study aimed to clarify the photosynthetic and storage functions of overwintering leaves in this species. METHODS: A 2-year experiment with defoliation and shading of overwintering leaves was conducted. Photosynthetic light response was measured in early spring (for overwintering leaves) and summer (for current-year leaves). KEY RESULTS: No nitrogen limitation of growth was detected in plants subjected to defoliation. The number of leaves, their size, reproductive activity (production of sori) and total leaf mass were not affected by the treatment. The defoliation of overwintering leaves significantly reduced the bulk density of rhizomes and the root weight. The carbohydrates consumed by the rhizomes were assumed to be translocated for leaf production. Photosynthetic products of overwintering leaves were estimated to be small. CONCLUSION: Overwintering leaves served very little as nutrient-storage and photosynthetic organs. They partly functioned as a carbon-storage organ but by contrast to previous studies, their physiological contribution to growth was found to be modest, probably because this species has a large rhizome system. The small contribution of overwintering leaves during the short-term period of this study may be explained by the significant storage ability of rhizomes in this long-living species. Other ecological functions of overwintering leaves, such as suppression of neighbouring plants in spring, are suggested.     

Why does Viola hondoensis (Violaceae) shed its winter leaves in spring?

? Premise of the study: Viola hondoensis is a perennial herb that inhabits the understory of temperate, deciduous forests. It is an evergreen plant with a leaf life span that is shorter than a year. Its summer leaves are produced in spring and shed in autumn; winter leaves are produced in autumn and shed in spring. Here we asked why the plant sheds its winter leaves in spring, though climate conditions improve from spring to summer. We proposed four hypotheses for the cause of shedding: (1) changes in seasonal environment such as day length or air temperature, (2) shading by canopy deciduous trees, (3) self-shading by taller summer leaves, and (4) competition for nutrients between summer and winter leaves. ? Methods: To test these hypotheses, we manipulated the environment of winter leaves: (1) plants were transplanted to the open site where there was no shading by canopy trees. (2) Petioles of summer leaves were anchored to the soil surface to avoid shading of winter leaves. (3) Sink organs were removed to eliminate nutrient competition. ? Key results: Longevity of winter leaves was extended when shading by summer leaves was eliminated and when sink organs were removed, but not when plants were transplanted to the open site. ? Conclusion: We conclude that the relative difference in light availability between summer and winter leaves is a critical factor for regulation of leaf shedding, consistent with the theory of maximization of the whole-plant photosynthesis.     

Seasonal pattern of photosynthetic production in a subalpine evergreen herb,<Emphasis Type="Italic"> Pyrola incarnata</Emphasis>

The seasonal pattern of growth and matter production of Pyrola incarnata, an evergreen herb on the forest floor in subalpine deciduous forests, was analyzed to understand the ecological significance of evergreenness in a subalpine climate with a short growing season and low temperature. Net production was highest under favorable light conditions in spring after the disappearance of snow cover, and 68% of the annual net production was attained before the canopy tree foliage had fully expanded. Most of the photosynthetic production in this period was carried out with over-wintered leaves. This appears to be an advantage of evergreenness. New leaves and inflorescences had developed in the period. Positive net production was maintained under deteriorating light conditions during summer, when 32% of the annual net production occurred. This production was used mainly for growth of fruits and underground organs. The net production of P. incarnata during summer was much higher than that of a related species that inhabits warm-temperate regions, because of its higher photosynthetic activity rather than its lower respiratory losses. The storage of dry matter in leaves and underground organs was not conspicuous. Unlike the warm-temperate species and another subalpine species that inhabits higher altitudes, P. incarnata is not strongly dependent on its reserve matter for the development of new organs.