Abstract Background and Aims The stomatal conductance (gs) of most plant species decreases in response to elevated atmospheric CO2 concentration. This response could have a significant impact on plant water use in a future climate. However, the regulation of the CO2-induced stomatal closure response is not fully understood. Moreover, the potential genetic links between short-term (within minutes to hours) and long-term (within weeks to months) responses of gs to increased atmospheric CO2 have not been explored. Methods We used Arabidopsis thaliana recombinant inbred lines originating from accessions Col-0 (strong CO2 response) and C24 (weak CO2 response) to study short- and long-term controls of gs. Quantitative trait locus (QTL) mapping was used to identify loci controlling short- and long-term gs responses to elevated CO2, as well as other stomata-related traits. Key Results Short- and long-term stomatal responses to elevated CO2 were significantly correlated. Both short- and long-term responses were associated with a QTL at the end of chromosome 2. The location of this QTL was confirmed using near-isogenic lines and it was fine-mapped to a 410-kb region. The QTL did not correspond to any known gene involved in stomatal closure and had no effect on the responsiveness to abscisic acid. Additionally, we identified numerous other loci associated with stomatal regulation. Conclusions We identified and confirmed the effect of a strong QTL corresponding to a yet unknown regulator of stomatal closure in response to elevated CO2 concentration. The correlation between short- and long-term stomatal CO2 responses and the genetic link between these traits highlight the importance of understanding guard cell CO2 signalling to predict and manipulate plant water use in a world with increasing atmospheric CO2 concentration. This study demonstrates the power of using natural variation to unravel the genetic regulation of complex traits.
We asked young scientists these questions: Have you ever encountered a particularly stark dif erence between an old and new position in your education or career? What was the dif erence between the positions, and what advice would you give to someone making a similar transition? Here, respondents share the challenges they faced when they took on new responsibilities and roles, changed fi elds, or moved to new places. To others in similar situations, they advise: Be confi dent, prepared, and patient; communicate; and always ask
Physical defenses and grazer avoidance of the bloom-forming microalga Gonyostomum semen may reduce the direct coupling between phytoplankton and higher trophic levels and result in an increased importance of alternative basal food resources such as bacteria and heterotrophic protozoans. To assess the importance of algal and heterotrophic food resources for zooplankton during G. semen blooms and the effects of zooplankton diets on a higher consumer, we analyzed the fatty acid composition of zooplankton and the invertebrate predator Chaoborus flavicans from eight lakes along a gradient in the predominance of G. semen relative to other algae and the duration of G. semen blooms. The proportion of fatty acids of bacterial origin increased significantly along the G. semen gradient in all consumers studied. In addition, the proportion of polyunsaturated fatty acids (PUFA) decreased in cladocerans. These results suggest that heterotrophic pathways can compensate for a reduced trophic coupling between phytoplankton and filter-feeding zooplankton. The lower PUFA content in cladoceran prey from lakes at the higher end of the G. semen gradient did not affect the PUFA content of the predator C. flavicans, suggesting selective assimilation and retention of PUFA and/or feeding on other, more PUFA-rich prey.
A number of marine and freshwater harmful algal bloom (HAB) species have colonized new areas and expanded their habitat range in recent years. Nevertheless it is notoriously difficult to establish when colonization first occurred, what the dispersal routes are, and to separate recent invasion from increases in existent but small populations. The freshwater raphidophyte Gonyostomum semen is a nuisance species that has expanded its habitat range and increased in abundance in northern Europe during the past decades. To evaluate to what extent sediments can be used for determining historic occurrence of G. semen, a quantitative real-time PCR method for detecting cysts of this algae was developed. This paper presents a qPCR protocol with a set of primers that are specific to Gonyostomum and with PCR conditions optimized for sediment samples from humic lakes, which are the common habitat of G. semen. With this sensitive method as few as 1.6 cysts per PCR reaction could be reliably quantified, corresponding to 320 cysts per g wet weight sediment. Cysts were present in sediments with ages ranging from years to decades and their persistence allows detection of historic populations up to at least 50 years old. With this qPCR assay it will be possible to trace the presence of G. semen in environments prior to the onset of algae-specific monitoring programs as well as for quantification in water column samples.
Stomata are small pores in the surface of plant leaves, balancing the uptake of CO2 against the loss of water vapour. As drought stress is projected to increase in many parts of the world, an improved understanding of how plants regulate their stomata in response to environmental stimuli may have important implications for securing food production in a future climate. The stomata of most plants respond to elevated CO2 concentration by partial closing, but the magnitude of this response shows a large variation among species and ecotypes. Plants that substantially decrease stomatal conductance under elevated CO2 reduce their water consumption and increase their water-use efficiency. However, the genetic basis for inter- and intraspecific variation in responsiveness is poorly understood, as are the mechanisms for sensing and responding to CO2. In the current study, we have measured the short-term stomatal response to elevated CO2 in a population of Arabidopsis thaliana recombinant inbred lines (RILs) originating from a cross between two parental genotypes showing a large difference in stomatal responsiveness. Hence, this RIL population showed a broad range in stomatal responsiveness, ranging from a 10% to a 60% reduction of stomatal conductance following an increase in CO2 concentration from 400 to 800ppm. Using quantitative trait locus (QTL) mapping we have identified two genetic loci involved in the stomatal response to elevated CO2. Potential candidate genes regulating the stomatal response will be discussed.
Small seedlings may be cheaper and establish faster than larger seedlings, but be more sensitive to stress and therefore less competitive. Site factors and species may also influence seedling performance, but the early performance of the most common species in Swedish plantations, Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) H. Karst), has rarely been compared at identical sites. To identify optimal species and size combinations, seedlings were grown in containers of two sizes, 50 and 90 cm(3), and planted during two consecutive years at both coastal and inland sites in northern Sweden. The larger Scots pine seedlings grew more rapidly than the smaller and were also the tallest of all seedlings five years after planting. No seedling size-related difference in Norway spruce growth was found. Highest mortality occurred the first three years after planting and pine weevils were one of the major causes. Larger seedlings of both species had a slightly lower mortality compared to smaller. Five years after planting, the proportion of damaged seedlings plateaued for Norway spruce, but not for Scots pine. Consequently, despite a higher growth the Scots pine seedlings remained sensitive to damage during a longer period than Norway spruce. No interactions were found between geographic location and seedling size or species on growth, mortality, or damage.
The large bloom-forming flagellate Gonyostomum semen has been hypothesized to be inedible to naturally occurring zooplankton due to its large cell size and ejection of long slimy threads (trichocysts) induced by physical stimulation. In a grazing experiment using radiolabelled algae and zooplankton collected from lakes with recurring blooms of G. semen and lakes that rarely experience blooms, we found that Eudiaptomus gracilis and Holopedium gibberum fed on G. semen at high rates, whereas Daphnia cristata and Ceriodaphnia spp. did not. Grazing rates of E. gracilis were similar between bloom-lakes and lakes with low biomass of G. semen, indicating that the ability to feed on G. semen was not a result of local adaptation. The high grazing rates of two of the taxa in our experiment imply that some of the nutrients and energy taken up by G. semen can be transferred directly to higher trophic levels, although the predominance of small cladocerans during blooms may limit the importance of G. semen as a food resource. Based on grazing rates and previous observations on abundances of E. gracilis and H. gibberum, we conclude that there is a potential for grazer control of G. semen and discuss why blooms of G. semen still occur.
Here, we use a novel space-by-time approach to study large-scale changes in phytoplankton species distribution in Swedish boreal lakes in response to climate variability. Using phytoplankton samples from 27 lakes, evenly distributed across Sweden, all relatively unimpacted by anthropogenic disturbance and sampled annually between 1996 and 2010, we found significant shifts in the geographical distribution of 18 species. We also found significant changes in the prevalence of 45 species (33 became more common and 12 less common) over the study period. Using species distribution models and phytoplankton samples from 60 lakes sampled at least twice between 1992 and 2010, we evaluated the importance of climate variability and other environmental variables on species distribution. We found that temperature (e.g., extreme events and the duration of the growing season) was the most important predictor for species detections. Many cyanobacteria, chlorophytes, and, to a lesser extent, diatoms and zygnematophytes, showed congruent and positive responses to temperature. In contrast, precipitation explained little variation and was important only for a few taxa (e.g., Staurodesmus spp., Trachelomonas volvocina). At the community level, our results suggest a change in community composition at temperatures over 20 °C and growing seasons longer than 40 days. We conclude that climate is an important driver of the distributional patterns of individual phytoplankton species and may drive changes in community composition in minimally disturbed boreal lakes.
Scarification is the most common measure to improve the planting environment in Sweden. However, different scarification methods give varying results. During the early 1990s, a nation-wide experiment with 10 field installations was established in order to test the effect of several scarification methods, including two intensities of soil inversion and mounding, on growth of planted Norway spruce seedlings and in comparison with no scarification (i.e. control). Eighteen growing seasons after planting, a higher seedling survival was found following soil inversion (77 per cent for normal and 76 per cent for intensive) compared with mounding (67 per cent) and control (57 per cent). The mean height of the planted trees across all sites 18 years after planting was 413 and 430 cm following normal and intensive soil inversion, respectively, 424 cm after mounding and 346 cm in the control. The difference in height between the scarification treatments and the control corresponded to a time gain of approximate to 4 years of growth after 18 years. However, the length of the leading shoot was not affected by scarification after 1418 years, indicating that scarification did not affect growth beyond the establishment phase. Scarification reduced variation in height of the planted trees. On scarified plots, the number of naturally regenerated trees increased with more than 100 per cent to reach a mean value of 2300 stems per hectare.
•We examine changes in distribution, occurrence and bloom incidence of the noxious Gonyostomum semen.•G. semen was spreading in southern Sweden but not north of Limes Norrlandicus.•The occurrence of G. semen was mainly explained by climate variables (e.g. temperature).•pH and water color were the best predictors of bloom incidence.•Morphometry and catchment variables were only important at high dominance levels of G. semen (>90% total phytoplankton biomass).
Blooms of the nontoxic raphidophyte Gonyostomum semen have shown a recent increase in frequency and distribution in the Fennoscandian region. Due to large cell size and several grazer-avoidance strategies, G. semen is hypothesized to be inedible for most zooplankton species and therefore may constitute a bottleneck for the transfer of energy and nutrients in pelagic food webs. Repression of other phytoplankton through increased competition and induced mortality could further exacerbate this effect. In a field study of four lakes with recurring blooms of G. semen and four lakes without blooms, we found significant differences in community structure between the two lake groups during the bloom period. Bloom-lakes had lower biovolumes of small chrysophytes and chlorophytes and zooplankton assemblages were predominated by small, potentially bacterivorous cladocerans, suggesting a limited availability of edible phytoplankton and an increased importance of microbial pathways during G. semen blooms. Low biovolumes of large cladocerans in bloom-lakes may be due to interference of G. semen with filter feeding. Moreover, high abundances of the phantom midge Chaoborus flavicans in bloom-lakes suggest that the flow of energy and nutrients is directed more towards this invertebrate predator than fish. This could have negative impacts on fish populations, especially if bloom periods are prolonged.
Fertilization and scarification are both performed to increase tree production at different stages of forest rotation periods. In this study, the effects of previous nitrogen fertilizations and scarification after clear felling on planted Pinus sylvestris L. seedlings and ground vegetation were investigated. Two fertilization experiments established around 1980 were harvested in 2006, after which the plots were scarified by disc trenching and re-planted. The plots had been repeatedly fertilized over a 20-year period before harvesting, with total N doses of 0, 450, 900 or 1800 kg N ha -1. After five growing seasons, the growth, survival and nutrient contents of the seedlings were measured, and ground vegetation was collected to estimate its biomass and nutrient content. Pre-harvest fertilization alone had only minor effects on the results, but scarification increased both the survival and growth of the planted seedlings. However, without scarification, seedling mortality increased with increasing fertilization intensity. The ground vegetation biomass was higher in plots without scarification, but the total biomass of seedlings and ground vegetation was similar in all treatments. Scarification thus favored seedling growth at the expense of ground vegetation. Only a few effects on nutrient content were found, but there were no signs of nutrient imbalance in any of the treatments. At higher levels of fertilization, the K:N ratio in the seedlings decreased while the K content in the ground vegetation increased. Overall, scarification had a greater impact than pre-harvest fertilization on the planted seedlings and the ground vegetation.
There is increasing evidence that forests and forest soil contribute to the signature of chlorine composition in water bodies. However, little is known about the potential effects of land management activities on chlorine biogeochemistry. This study examines the effects of previous nitrogen addition on chlorine chemistry in a Pinus sylvestris L. forest located in south-central Sweden (60°00′N, 13°43′E). Repeated addition of nitrogen to study plots over a 20-year period resulted in total additions of 0, 450 and 900 kg N ha −1 . Soil samples were collected before harvesting, and soil solution and biomass were sampled following final felling. Contrary to previous findings, we found no clear evidence that previous addition of nitrogen had hampered the formation of organic chlorine in the organic soil layer. We suggest that the effects of nitrogen addition on chlorination processes are not seen in the surface soil, but are instead manifested in the migration of organic matter in the mineral soil. Soil organic matter from the E-horizon had a lower chlorination degree in the nitrogen-amended plots than in the control plots. In addition, we observed lower Cl − levels in the seedling needles following high nitrogen fertilization (900 N) than in 450 N or control. These results add on the importance of studying chlorine dynamics with focus on what chlorinated soil organic matter can be resistant to degradation compared to average soil organic matter and what can be leached as a result of harvesting and available for vegetation.
The purpose of the study was to create a near optimal environment for seedling establishment and growth, without the restrain of water and nutrients but under climate conditions typical for the region. This to give us valuable knowledge about the growth potential of different seedling types in the field. The experimental site was situated in southern Sweden. Six treatment combinations were applied including two site treatments; 1) soil inversion, i.e. the control treatment, and 2) soil inversion, drip irrigation and fertilization combined with plastic cover mulch, i.e. the optimization treatment, and three seedling types of Norway spruce (Picea abies L. Karst.), (a) a 2-year-old Plug+1 seedling, (b) a 1.5-year-old containerized seedling and (c) a 10-week-old mini seedling. Effects on seedling nutrient status and growth were studied during the first three years after planting. Height, diameter and biomass of the seedlings grown in the optimized environment were significantly greater than for seedlings grown in the control. The Plug+1 seedlings grown in the optimization treatment had, after three years, reached a height of 124 cm, while the containerized seedlings were 104 cm and the mini seedlings 45 cm. In practical plantations, this height is usually gained after 5â10 years depending on planting conditions. Biomass partitioning did not differ between optimization treatments, but between seedling types. The mini seedlings allocated less biomass to the roots and more biomass to needles and stem in comparison with the two other seedling types. Mini seedlings also broke bud earlier. Throughout the experimental period, seedling nutrient status for all treatment combinations was followed and a balanced nutrient supply of macro- and micronutrients was given in the optimization treatment. Nutrient concentrations were constantly higher in seedlings grown in the optimization treatment, but the difference decreased over time. Results from this study shows that, by improving site conditions associated with fast establishment, growth check can be avoided.
Successful regeneration of conifer forests by planting is, in large parts of Europe, highly dependent on the effective suppression of damage caused by the pine weevil Hylobius abietis. We investigated the effectiveness of various combinations of control measures against pine weevil damage under boreal forest conditions in Sweden. In particular, we aimed to determine whether satisfactory regeneration could be obtained without the use of insecticides. The experimental study was established on ten new clear-cuts in each of three consecutive years. We studied the use of chemical and physical methods to protect seedlings directly, and investigated the influence of seedling type, age of clear-cut, and a number of soil factors as affected by preparation of the planting site, on the mortality and level of damage suffered by protected and unprotected seedlings. After two seasons, mortality due to pine weevil was 16% among unprotected seedlings, 6% for seedlings treated with the insecticides cypermethrin or imidacloprid, and less than 1% for those physically protected by a coating of Conniflex. However, the Conniflex, which consists of fine-grained sand embedded in a flexible acrylate matrix, was applied manually, and this may have enhanced its effectiveness compared to that achieved during large-scale, commercial application. Two types of containerized Norway spruce seedlings, which differed mainly in their stem diameter (average 2.6 mm and 3.5 mm), were used in the experiments. Among the unprotected seedlings, the narrower stemmed type was more frequently attacked (34% vs. 28%) and killed (19% vs. 12%) by pine weevil. Mortality caused by pine weevil among unprotected seedlings was higher on 1-year-old than on 2-year-old clear-cuts (20% vs. 12%). Soil preparation around unprotected seedlings had a substantial effect on the proportion attacked and killed by pine weevil as well as on the total mortality, with the highest level of feeding damage and mortality occurring on seedlings in undisturbed humus, and the lowest levels occurring on seedlings planted in pure mineral soil (26% vs. 7% for unprotected seedlings). This study demonstrates that acceptable levels of seedling survival can be achieved in regenerations of North European boreal forest without the use of insecticides. Mortality of unprotected seedlings can be reduced to acceptable levels if they are mostly planted in pure mineral soil. Damage can be further reduced by using seedlings with a somewhat larger stem diameter. Insecticides or a physical barrier of Conniflex alone appear to provide a sufficient level of protection. (C) 2011 Elsevier B.V. All rights reserved.
Interactions between scarification treatments and seedling types of Norway spruce were examined at two different locations in southern Sweden. The scarification treatments were not scarified control, mounding and soil inversion and the seedling types were a 10 week-old containerized seedling (mini seedling), a 2 year-old containerized seedling and a 2 year-old hybrid seedling, grown as a containerized seedling the first year and as a bare-root seedling the second year. Site preparation intensity had a greater effect on the mini seedlings. The results indicated that mini seedlings established faster in the soil inversion treatment compared to the larger seedling types. At the same seedling age, the mini seedlings had faster or similar growth rate as the containerized and the hybrid seedlings regardless of scarification treatment. Although mini seedlings grew as well as or even better than larger seedlings if they were successfully established, they were more sensitive to their planting environment and proper handling was critical. Problems with frost heaving and competing vegetation have to be taken into consideration when choosing site preparation method and seedling type.
Establishment of Norway spruce (Picea abies (L.) Karst.) seedlings is often restricted by competition from vegetation, drought, and damage by pine weevils. In this study, effects of mulching on these factors were investigated. Norway spruce seedlings were planted on fresh and 1-year-old clearcuts treated with mulch on three sites in southern Sweden. Mulch was made of slash from the old stand and applied on whole blocks at three different depths: 0, 10, and 20 cm. Both insecticide-treated and untreated seedlings were planted. By reducing the competing vegetation and improving soil moisture and mineralization, mulching created a favorable growth environment. Mulching significantly improved growth in terms of height, diameter, and volume of the seedlings. Growth continued to increase over time in mulched treatments, probably as an effect of increased nutrient availability. The 20 cm mulch layer generated the greatest increase in growth throughout the 10-year experimental period. Soil moisture was preserved under the isolating mulch layer and during periods of drought soil water potential was significantly higher in mulched treatments. After 2 years, percent cover of competing vegetation was 50%–60% without mulch and 10%–20% with a mulch depth of 20 cm. Insecticide-treated seedlings achieved a survival rate close to 100% in all mulching treatments, whereas survival among untreated seedlings was only 40% on some clearcuts. Mulching alone did not affect survival or abundance of pine weevils.
The effects of mulch and scarification on the establishment and growth of six clones of Norway spruce [ Picea abies ( L.) Karst.] were examined in a study in southern Sweden. Mulch and scarification reduced the amount of competing vegetation and tended to improve soil moisture and soil temperature. Budbreak occurred earlier for cuttings planted in the scarification treatment in comparison with the control and mulching treatment. Scarification increased survival and the combination of scarification and mulch resulted in the best growth. Gas exchange and the number of new roots were higher in planting spots covered with mulch. Clonal differences regarding gas exchange and growth were significant. Clones with a poor height growth also had a low gas exchange and a small number of new roots. In contrast, the clone with the best height growth had high gas exchange and a large number of new roots. Clonal effects on growth were greater than site preparation effects.