Intermittent low temperatures constrain spring recovery of photosynthesis in boreal Scots pine forests

During winter and early spring, evergreen boreal conifers are severely stressed because light energy cannot be used when photosynthesis is pre‐empted by low ambient temperatures. To study photosynthetic performance dynamics in a severe boreal climate, seasonal changes in photosynthetic pigments, chloroplast proteins and photochemical efficiency were studied in a Scots pine forest near Zotino, Central Siberia. In winter, downregulation of photosynthesis involved loss of chlorophylls, a twofold increase in xanthophyll cycle pigments and sustained high levels of the light stress‐induced zeaxanthin pigment. The highest levels of xanthophylls and zeaxanthin did not occur during the coldest winter period, but rather in April when light was increasing, indicating an increased capacity for thermal dissipation of excitation energy at that time. Concomitantly, in early spring the D1 protein of the photosystem II (PSII) reaction centre and the light‐harvesting complex of PSII dropped to their lowest annual levels. In April and May, recovery of PSII activity, chloroplast protein synthesis and rearrangements of pigments were observed as air temperatures increased above 0°C. Nevertheless, severe intermittent low‐temperature episodes during this period not only halted but actually reversed the physiological recovery. During these spring low‐temperature episodes, protective processes involved a complementary function of the PsbS and early light‐induced protein thylakoid proteins. Full recovery of photosynthesis did not occur until the end of May. Our results show that even after winter cold hardening, photosynthetic activity in evergreens responds opportunistically to environmental change throughout the cold season. Therefore, climate change effects potentially improve the sink capacity of boreal forests for atmospheric carbon. However, earlier photosynthesis in spring in response to warmer temperatures is strongly constrained by environmental variation, counteracting the positive effects of an early recovery process.     

Above-ground biomass and structure of pristine Siberian Scots pine forests as controlled by competition and fire

The study presents a data set of above-ground biomass (AGB), structure, spacing and fire regime, for 24 stands of pristine Siberian Scots pine (Pinus sylvestris) forests with lichens (n = 20) or Vaccinium/mosses (n = 4) as ground cover, along four chronosequences. The stands of the “lichen” site type (LT) were stratified into three chronosequences according to stand density and fire history. Allometric equations were established from 90 sample trees for stem, coarse branch, fine branch, twig and needle biomass. The LT stands exhibited a low but sustained biomass accumulation until a stand age of 383 years. AGB reached only 6–10 kg_dw m⁻² after 200 years depending on stand density and fire history compared to 20 kg_dw m⁻² in the “Vaccinium” type (VT) stands. Leaf area index (LAI) in the LT stands remained at 0.5–1.5 and crown cover was 30–60%, whereas LAI reached 2.5 and crown cover was >100% in the VT stands. Although nearest-neighbour analyses suggested the existence of density-dependent mortality, fire impact turned out to have a much stronger effect on density dynamics. Fire scar dating and calculation of mean and initial fire return intervals revealed that within the LT stands differences in structure and biomass were related to the severity of fire regimes, which in turn was related to the degree of landscape fragmentation by wetlands. Self-thinning analysis was used to define the local carrying capacity for biomass. A series of undisturbed LT stands was used to characterise the upper self-thinning boundary. Stands that had experienced a moderate fire regime were positioned well below the self-thinning boundary in a distinct fire-thinning band of reduced major axis regression slope −0.26. We discuss how this downward shift resulted from alternating phases of density reduction by fire and subsequent regrowth. We conclude that biomass in Siberian Scots pine forests is strongly influenced by fire and that climate change will affect ecosystem functions predominantly via changes in fire regimes. Received: 2 July 1998 / Accepted: 10 June 1999     

Seed mass versus seedling performance in Scots pine: a maternally dependent trait

It is generally accepted that larger seeds give rise to seedlings with better performance. On the other hand, the size that a seed reaches is genetically determined by at least two different traits ; the genetic variability of the developing embryo and the genetic variability of the maternal plant. Thus, the relative contributions of these two traits affect seedling performance by influencing seed size. In this paper, I investigate the effect of seed size on seedling performance in the Scots pine ( Pinus sylvestris ). From eight maternal plants, 50 seeds were planted in each of two soil types (800 seeds in total), and seedling performance was monitored for 1 yr. Seed mass proved to be highly constant within maternal plants. Soil type influenced emergence and survival; however, the effect of soil type differed depending on maternal origin. Seed mass was positively correlated with seedling emergence, although this relationship was not found for seedling survival or date of emergence. The initial growth of the shoot was also positively correlated with seed mass. However, after one growing season, seed mass had no effect on seedling performance, which depended exclusively on maternal origin. Nevertheless, the mean mass of seeds produced by plants was positively correlated with mean values of growth parameters. Thus, first-year seedling performance seems to be a maternal trait indirectly associated with seed size.     

Xylem cavitation in two mature Scots pine forests growing in a wet and a dry area of Britain

The xylem cavitation rate, stem water content, stomatal conductance and leaf water potential of mature Scots pine (Pinus sylvestris L.) were compared over a period of 18 months at two sites in Britain where trees were planted from the same seed source. The sites were at Thetford in south-east England, where the climate is relatively warm and dry (average rainfall of 600mm per year), and Aberfoyle in central Scotland, where it is relatively cool and wet (average rainfall of 1500 mm year^?1). In the first year of study (1992) the natural difference between the sites was amplified by a severe drought affecting south-east England. Acoustic emissions (as a result of cavitation) were detected at both sites, but were not an everyday occurrence, and rates depended on short-term meteorological variation. The relative water content (RWC) of the trunk at breast height at Thetford was significantly lower than at Aberfoyle, and declined in parallel with increasing severity of drought. Following the release from drought a gradual increase in RWC was found, but the pre-drought RWC was not attained. The same xylem water potential gradient and stomatal conductance was found at the two sites; but on a diurnal basis, as water potential declined, cavitation rates increased. There was no difference in vulnerability to cavitation or in hydraulic conductance between the sites.     

Ribulose bisphosphate carboxylase, protein and nitrogen in Scots pine seedlings cultivated at different nutrient levels

Seedlings of Scots pine (Pinus sylvestris L.) of a northern provenance were cultivated in nutrient solution for 10 weeks in a climate chamber. The nutrient solution (renewed by solution exchange) contained 2.5, 10 or 50 mg N I^?1. All other essential elements were added in optimal proportion to the nitrogen. Seedlings cultivated at 10 and 50 mg N I^?1 were similar with respect to all characteristics studied. Seedlings cultivated at 2.5 mg N I^?1 showed a lower growth rate, especially for the shoot, and an altered morphology, with high root:shoot ratios and long, slender roots. The nitrogen concentrations in shoot and needles as well as in whole seedlings were not significantly affected by the nitrogen supply, while the nitrogen concentrations in the roots were somewhat lower at 2.5 mg N I^?1. Ribulose bisphosphate carboxylase (EC 4.1.1.39) activity and the concentrations of carboxylase, total and soluble protein and of chlorophyll in the needles were consistently much lower for seedlings cultivated at 2.5 mg N I^?1, than for seedlings grown at higher nutrient levels. A close correlation was observed between activity and concentration of the carboxylase (r=0.95). Carboxylase activity and protein were more sensitive to a low nutrient supply than was chlorophyll. The data show how activity and concentration of ribulose bisphosphate carboxylase and the concentrations of soluble and total protein and of chlorophyll in needles of pine seedlings can be negatively affected by the nutrient supply, also when the nitrogen concentrations in the needles are close to those observed at optimal nutrient supply. It is suggested that pine seedlings store assimilated non-protein nitrogen in the needles when protein synthesis is under restraint. The nitrogen concentration in needles and seedlings could not be used as a measure of the physiological state of the seedlings.     

Seed colour and post-fire bird predation in a Mediterranean pine forest

In a Pinus halepensis Mill. forest, a field experiment was designed to evaluate post-fire seed predation as affected by combinations of seed colour and soil substrates: light grey and black seeds combined with light grey ash, dark grey ash and pale brown sand. A survey of bird species inhabiting the area was also carried out and polyphenolic content of seed coat was assessed in seed lots of different colour. Light grey seeds were observed to be less predated on light grey ash, suggesting eucrypsis as a protective strategy against bird predation. On the contrary, no clear pattern was observed for the predation of black seeds on different substrates. In the study area both bird species breaking the seed coat and eating the endosperm and bird species swallowing the whole seed were monitored. We have estimated that more seeds were swallowed than broken, in all colour categories. Light grey seeds, which were found to have a higher content of polyphenols, were predated more than black seeds when exposed on the same substrate. Thus, no evidence was produced that the amount of polyphenols in seed coat could protect seeds from predation.     

Inhibitors of endogenous proteinases in the seeds of Scots pine, Pinus sylvestris

Extracts of resting pine seeds inhibited the proteinase activities present in extracts of endosperms of germinating seeds (hydrolysis of haemoglobin at pH 3.7 and hydrolysis of casein at pH 5.4 and 7.0). Heating the extracts of resting seeds at 60°C destroyed their own proteinase activity but their proteinase inhibitor activity decreased by only 25 to 30%. Some properties of the inhibitor(s) were studied using extracts treated at 60°C. The inhibitor activities were non-dialysable. the inhibition increased linearly with increasing inhibitor concentration up to 80% of total proteinase activity, and the maximal inhibition was 80% at pH 3.7. 90% at pH 5.4. and 97% at pH 7.0. The extracts of resting seeds did not inhibit the pepsin-like acid pine proteinase that accounts for a minor part of the proteolytic activity of endosperm extracts at pH 3.7. Neither did they have any effect on the acid pine carboxypeptidase or trypsin and chymotrypsin. Fresh extracts of endosperms of germinating seeds contained relatively high proteinase activity (assayed directly) and moderate inhibitor activity (assayed after treatment at 60°C). When fresh extracts were dialysed at 50°C for 48 h their proteinase activities increased considerably while the corresponding inhibitor activities disappeared. It is concluded that the decrease of inhibitors during dialysis is due to enzymatic inactivation and that the corresponding increase of proteinase activities is at least partly due to the destruction of the inhibitors.     

Gas exchange in a 20-year-old stand of Scots pine

The rates of net photosynthesis and transpiration of one-year-old shoots were measured in situ in five different positions within the crown of a young Scots pine ( Pinus sylvestris L.). Measurements were carried out on south- and north-facing shoots on the third and sixth whorls, respectively, and on an east-facing shoot on the ninth whorl. In another investigation the rates of gas exchange of one-year-old shoots on the third whorl of eight different trees were studied. The measurements were made during June and July, 1977, under non-limiting conditions of soil water. The daily rates of net photosynthesis in whorls three and six followed the light conditions closely, with higher rates for the south side of each whorl and higher for whorl three than six. On whorl nine the shoot had a higher light compensation point and a low rate of photosynthesis at light saturation compared to the other shoot positions. The quantum yield for the shoot on the lowest whorl, as estimated from the linear part of the light response curve, was 50% lower than for shoots on whorl three and six.
The variation in transpiration rates was pronounced within the crown as an effect of differences in the absolute value and diurnal course of stomatal conductance. The variation in net photosynthesis was small between different trees while the variation in transpiration was much higher. Thus the variation in water use efficiency was great. It is concluded that it is possible to extrapolate measurements of net photosynthesis from individual trees up to a stand level without introducing large errors in the estimate. More caution must be paid before extrapolating tree transpiration up to stand transpiration. However, before an extrapolation of gas exchange is made from tree to stand level the variation in net photosynthesis and transpiration rate within the crown must be known.     

Resilience,vulnerability and conservation strategies in high-mountain pine forests in the Gredos Range,central Spain

Background: High-mountain pine forests and broom communities in central Spain today have led to contrasting interpretations of their natural or human-induced origin.

Aims: We evaluated the vulnerability and resilience of high-mountain Pinus sylvestris/P. nigra forests and derived Cytisus broom scrub communities to climate and anthropogenic disturbances.

Methods: We assessed historical transitions from forest to scrub and their relation to climate and human influences, using a multi-proxy paleoenvironmental study (pollen, non-pollen palynomorphs, charcoal, magnetic susceptibility) in three mires in the Gredos Range, western Iberian Central System.

Results: High percentages of Pinus sylvestris/nigra pollen and the identification of their macro remains demonstrated that high-mountain pine forests have been present in the oromediterranean bioclimatic belt of the Gredos Range since the mid-Holocene. After that, a major human-induced decline, enhanced by climate conditions, has led to their gradual replacement by broom communities.

Conclusions: Broom communities are derived from ancient pine forests that were intensively transformed by human activities after 700 cal year BP, and largely disappeared by ca. 500 cal yr BP. Today’s landscape, dominated by broom scrub and grasslands with scattered stands of pines, shows high resilience and provides suitable refugia for a rich mountain biodiversity which deserves a further protection.     

Fire and site type effects on the long-term carbon and nitrogen balance in pristine Siberian Scots pine forests

Effects of fire and site type on carbon (C) and nitrogen (N) balances were determined by following the change of total and component C and N pools along four chronosequences of fire-prone Siberian Scots pine ecosystems. These differed in the mean return interval of surface fires (unburned – moderately burned, 40 years – heavily burned, 25 years) and site quality (lichen versus Vaccinium site type). Of the Vaccinium site type (higher site quality) only a moderately burned chronosequence was studied. A total of 22 even-aged stands were investigated with stand ages ranging from 2 to 383 years. The C balance was dominated by the opposing dynamics of coarse woody debris (CWD) and biomass and could be divided into three phases: (1) Young stands (up to 40 years)acted as a net source for C of 6-10 mol C m-2 year-1 because the previous generation CWD pool originating from stand-replacing crown fires decayed much faster than biomass increased. During this period the C pool in the unburned lichen type chronosequence decreased from 807 to 480 mol C m-2. (2) Middle aged stands (40-100 years) being in a stage of maximum biomass accumulation were a net sink of 8-10 mol C m-2 year-1. (3)Maturestands (100 to > 350 years) continued to sequester C at a lower rate (0.8-2.5mol C m-2 year-1). Differences in the rates of C sequestration during the two later phases could be explained by the complex interaction between surface fire regime and site type. Recurrent surface fires resulted in enhanced mortality and regularly redistributed C from the living to the CWD pool thereby lowering the rate of C sequestration. Site quality determined the potential to recover from disturbance by fire events. Differences in site type did not correlate with soil and total ecosystem N pool size. However, the N status of needles as well as the N pool of physiologically active tissue was highest in the stands of the Vaccinium type. The woody C pool (biomass + CWD) was sensitive to differences in surface fire regime and site type. It was lowest in the heavily burned lichen type chronosequence (297 ± 108 mol C m-2), intermediate in the unburned and moderately burned lichen type chronosequence (571 ± 179 mol C m-2) and highest in the moderately burned Vaccinium type chronosequence (810 ± 334 mol C m-2). In contrast, the total soil C pool (organic plus mineral layer down to a depth of 25 cm) was independent of stand age, surface fire regimeand site type and fluctuated around a value of 250 mol C m-2. The organic layer C pool oscillated in response to recurring surface fires and its C pool was dependent on time since fire increasing at a rate of about 1.5 mol C m-2 year-during the first 40 years and then reaching a plateau of 170 mol C m-2. The total ecosystem N pool was 7.4 ± 1.5 mol N m-2 on average of which only 25 % were stored in biomass or coarse woody debris. Total ecosystem N was independent of stand age, surface fire regime and site type. No correlation was found between total ecosystem C and N pools. Average total ecosystem C:N ratio was 114 ± 35 mol C mol N-1. A conceptual model illustrating how changes in the regime of stand-replacing crown fires and recurrent surface fires and changes in site quality interact in determining the long-term C balance in Siberian Scots pine forests is presented.