Enriching plant diversity in grasslands by large-scale experimental sward disturbance and seed addition along gradients of land-use intensity

Aims The relationship between biodiversity and ecosystem functioning is among the most active areas of ecological research. Furthermore, enhancing the diversity of degraded ecosystems is a major goal in applied restoration ecology. In grasslands, many species may be locally absent due to dispersal or microsite limitation and may therefore profit from mechanical disturbance of the resident vegetation. We established a seed addition and disturbance experiment across several grassland sites of different land use to test whether plant diversity can be increased in these grasslands. Additionally, the experiment will allow us testing the consequences of increased plant diversity for ecosystem processes and for the diversity of other taxa in real-world ecosystems. Here we present details of the experimental design and report results from the first vegetation survey one year after disturbance and seed addition. Moreover, we tested whether the effects of seed addition and disturbance varied among grassland depending on their land use or pre-disturbance plant diversity. Methods A full-factorial experiment was installed in 73 grasslands in three regions across Germany. Grasslands were under regular agricultural use, but varied in the type and the intensity of management, thereby representing the range of management typical for large parts of Central Europe. The disturbance treatment consisted of disturbing the top 10 cm of the sward using a rotavator or rotary harrow. Seed addition consisted of sowing a high-diversity seed mixture of regional plant species. These species were all regionally present, but often locally absent, depending on the resident vegetation composition and richness of each grassland. Important findings One year after sward disturbance it had significantly increased cover of bare soil, seedling species richness and numbers of seedlings. Seed addition had increased plant species richness, but only in combination with sward disturbance. The increase in species richness, when both seed addition and disturbance was applied, was higher at high land-use intensity and low resident diversity. Thus, we show that at least the early recruitment of many species is possible also at high land-use intensity, indicating the potential to restore and enhance biodiversity of species-poor agricultural grasslands. Our newly established experiment provides a unique platform for broad-scale research on the land-use dependence of future trajectories of vegetation diversity and composition and their effects on ecosystem functioning.

Moderately High Temperatures Inhibit Ribulose-1,5-Bisphosphate Carboxylase/Oxygenase (Rubisco) Activase-Mediated Activation of Rubisco

We tested the hypothesis that light activation of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) is inhibited by moderately elevated temperature through an effect on Rubisco activase. When cotton (Gossypium hirsutum L.) or wheat (Triticum aestivum L.) leaf tissue was exposed to increasing temperatures in the light, activation of Rubisco was inhibited above 35 and 30°C, respectively, and the relative inhibition was greater for wheat than for cotton. The temperature-induced inhibition of Rubisco activation was fully reversible at temperatures below 40°C. In contrast to activation state, total Rubisco activity was not affected by temperatures as high as 45°C. Nonphotochemical fluorescence quenching increased at temperatures that inhibited Rubisco activation, consistent with inhibition of Calvin cycle activity. Initial and maximal chlorophyll fluorescence were not significantly altered until temperatures exceeded 40°C. Thus, electron transport, as measured by Chl fluorescence, appeared to be more stable to moderately elevated temperatures than Rubisco activation. Western-blot analysis revealed the formation of high-molecular-weight aggregates of activase at temperatures above 40°C for both wheat and cotton when inhibition of Rubisco activation was irreversible. Physical perturbation of other soluble stromal enzymes, including Rubisco, phosphoribulokinase, and glutamine synthetase, was not detected at the elevated temperatures. Our evidence indicates that moderately elevated temperatures inhibit light activation of Rubisco via a direct effect on Rubisco activase.

Knock-out of Arabidopsis metal transporter gene IRT1 results in iron deficiency accompanied by cell differentiation defects

IRT1 and IRT2 are members of the Arabidopsis ZIP metal transporter family that are specifically induced by iron deprivation in roots and act as heterologous suppressors of yeast mutations inhibiting iron and zinc uptake. Although IRT1 and IRT2 are thought to perform redundant functions as root-specific metal transporters, insertional inactivation of the IRT1 gene alone results in typical symptoms of iron deficiency causing severe leaf chlorosis and lethality in soil. The irt1 mutation is characterized by specific developmental defects, including a drastic reduction of chloroplast thylakoid stacking into grana and lack of palisade parenchyma differentiation in leaves, reduced number of vascular bundles in stems, and irregular patterns of enlarged endodermal and cortex cells in roots. Pulse labeling with 59Fe through the root system shows that the irt1 mutation reduces iron accumulation in the shoots. Short-term labeling with 65Zn reveals no alteration in spatial distribution of zinc, but indicates a lower level of zinc accumulation. In comparison to wild-type, the irt1 mutant responds to iron and zinc deprivation by altered expression of certain zinc and iron transporter genes, which results in the activation of ZIP1 in shoots, reduction of ZIP2 transcript levels in roots, and enhanced expression of IRT2 in roots. These data support the conclusion that IRT1 is an essential metal transporter required for proper development and regulation of iron and zinc homeostasis in Arabidopsis.

Effect of locally increased zinc contents on zinc transport from the flag leaf lamina to the maturing grains of wheat

From previous experiments, it was evident that the accumulation of zinc in maturing wheat grains is highly regulated, but the regulatory mechanisms involved are not yet identified. In this study, we determined the transfer of radiolabelled zinc (fed directly into a leaf flap) from the flag leaf lamina to the grains. We also determined how this zinc transfer was affected by feeding additional unlabeled zinc (1 μmol per plant) either into the flag leaf sheath or the peduncle. Most of the 65Zn was retained in the feeding flap. A high percentage of the zinc exported from the flap accumulated in the grains with little accumulation of radiolabel in the other parts of the shoot. Unlabeled zinc remained mainly in the feeding flap and in the parts reached by the transpiration stream from the feeding position. The transfer of radiolabelled zinc was essentially not influenced by unlabeled zinc fed into another plant part. Our results suggest that the loading of zinc into the phloem and the mass flow in the sieve tubes might regulate zinc redistribution within the wheat shoot.

Long-term Responses of Mountain Ecosystems to Environmental Changes: Resilience, Adjustment, and Vulnerability

The steep environmental gradients of mountain ecosystems over short distances reflect large gradients of several climatic parameters and hence provide excellent possibilities for ecological research on the effects of environmental change. To gain a better understanding of the dynamics of abiotic and biotic parameters of mountain ecosystems, long-term records are required since permanent plots in mountain regions cover in the best case about 50 - 70 years. In order to extend investigations of ecological dynamics beyond these temporal limitations of permanent plots, paleoecological approaches can be used if the sampling resolution can be adapted to ecological research questions, e.g. a sample every 10 years. Paleoecological studies in mountain ecosystems can provide new ecological insights through the combination of different spatial and temporal scales. [f we thus improve our understanding of processes across both steep environmental gradients and different time scales, we may be able to better estimate ecosystem responses to current and future environmental change (Ammann et al. 1993; Lotter et al. 1997). The complexity of ecological interactions in mountain regions forces us to concentrate on a number of sub-systems - without losing sight of the wider context. Here, we summarize a few case studies on the effects of Holocene climate change and disturbance on the vegetation of the Western Alps. To categorize the main response modes of vegetation to climatic change and disturbance in the Alps we use three classes of ecological behaviour: "resilience", "adjustment", and "vulnerability", We assume a resilient (or elastic) behaviour if vegetation is able to recover to its former state, regaining important ecosystem characteristics, such as floristic composition, biodiversity, species abundances, and biomass (e.g. Küttel 1990; Aber and Melillo 199 1). Conversely, vegetation displacements may occur in response to climatic change and/or disturbance. In some cases, this may culminate in irreversible large-scale processes such as species and/or community extinctions. Such drastic developments indicate high ecosystem vulnerability (or inelasticity or instability, for detailed definitions see Küttel 1990; Aber and Melillo 199 1) to climatic change and/or disturbance. In this sense, the "vulnerability" (or instability) of an ecosystem is expressed by the degree of failure to recover to the original state before disturbance and/or climatic change. Between these two extremes (resilience vs. vulnerability), ecosystem adjustments to climatic change and/or disturbance may occur, including the appearance of new and/or the disappearance of old species. The term "adjustment" is hence used to indicate the response of vegetational communities, which adapted to new environmental conditions without losing their main character. For forest ecosystems, we assume vegetational adjustments (rather than vulnerability) if the dominant (or co-dominant) tree species are not outnumbered or replaced by formerly unimportant plant species or new invaders. Adaptation as a genetic process is not discussed here and will require additional pbylogeographical studies (that incorporate the analysis of ancient DNA) in order to fully understand the distributions of ecotypes.

PLANT MACROFOSSIL METHODS AND STUDIES | Treeline Studies

Macrofossil series from mountainous and polar areas such as the Alps, the Scandes, and the Rocky Mountains provide useful information about the late Quaternary course of treelines. Reconstructions of past treeline positions and compositions after the Ice Age can be used to infer paleoclimatic information or to study treeline ecosystem responses to (rapid) climatic change. Both approaches are illustrated by study cases. In addition to these paleoclimatic and paleoecological issues, macrofossil records can be used to validate dynamic treeline vegetation models. An example from the Alps shows the high potential of such a procedure towards applying forest-succession models for assessing future vegetation dynamics at treeline.

Lake-effect snow as the dominant control of mesic-forest distribution in Michigan, USA

1Recent studies demonstrated the sensitivity of northern forest ecosystems to changes in the amount and duration of snow cover at annual to decadal time scales. However, the consequences of snowfall variability remain uncertain for ecological variables operating at longer time scales, especially the distributions of forest communities. 2The Great Lakes region of North America offers a unique setting to examine the long-term effects of variable snowfall on forest communities. Lake-effect snow produces a three-fold gradient in annual snowfall over tens of kilometres, and dramatic edaphic variations occur among landform types resulting from Quaternary glaciations. We tested the hypothesis that these factors interact to control the distributions of mesic (dominated by Acer saccharum, Tsuga canadensis and Fagus grandifolia) and xeric forests (dominated by Pinus and Quercus spp.) in northern Lower Michigan. 3We compiled pre-European-settlement vegetation data and overlaid these data with records of climate, water balance and soil, onto Landtype Association polygons in a geographical information system. We then used multivariate adaptive regression splines to model the abundance of mesic vegetation in relation to environmental controls. 4Snowfall is the most predictive among five variables retained by our model, and it affects model performance 29% more than soil texture, the second most important variable. The abundance of mesic trees is high on fine-textured soils regardless of snowfall, but it increases with snowfall on coarse-textured substrates. Lake-effect snowfall also determines the species composition within mesic forests. The weighted importance of A. saccharum is significantly greater than of T. canadensis or F. grandifolia within the lake-effect snowbelt, whereas T. canadensis is more plentiful outside the snowbelt. These patterns are probably driven by the influence of snowfall on soil moisture, nutrient availability and fire return intervals. 5Our results imply that a key factor dictating the spatio-temporal patterns of forest communities in the vast region around the Great Lakes is how the lake-effect snowfall regime responds to global change. Snowfall reductions will probably cause a major decrease in the abundance of ecologically and economically important species, such as A. saccharum.

Quaternary refugia of the sweet chestnut (Castanea sativa Mill.): an extended palynological approach

Knowledge about the glacial refugia of the thermophilous European Castanea sativa Mill. (sweet chestnut) is still inadequate. Its original range of distribution has been masked by strong human impact. Moreover, under natural conditions the species was probably admixed with other taxa (such as Quercus, Fraxinus, Fagus, Tilia) and thus possibly represented by low percentages in pollen records. In this paper we try to overcome the difficulties related to the scarcity and irregularity of chestnut pollen records by considering 1471 sites and extending the palynological approach to develop a Castanea refugium probability index (IRP), aimed at detecting possible chestnut refugia where chestnuts survived during the last glaciation. The results are in close agreement with the current literature on the refugia of other thermophilous European trees. The few divergences are most probably due to the large amount of new data integrated in this study, rather than to fundamental disagreements about data and data interpretation. The main chestnut refugia are located in the Transcaucasian region, north-western Anatolia, the hinterland of the Tyrrhenian coast from Liguria to Lazio along the Apennine range, the region around Lago di Monticchio (Monte Vulture) in southern Italy, and the Cantabrian coast on the Iberian peninsula. Despite the high likelihood of Castanea refugia in the Balkan Peninsula and north-eastern Italy (Colli Euganei, Monti Berici, Emilia-Romagna) as suggested by the IRP, additional palaeobotanical investigations are needed to assess whether these regions effectively sheltered chestnut during the last glaciation. Other regions, such as the Isère Département in France, the region across north-west Portugal and Galicia, and the hilly region along the Mediterranean coast of Syria and Lebanon were classified as areas of medium refugium probability. Our results reveal an unexpected spatial richness of potential Castanea refugia. It is likely that other European trees had similar distribution ranges during the last glaciation. It is thus conceivable that shelter zones with favourable microclimates were probably more numerous and more widely dispersed across Europe than so far assumed. In the future, more attention should be paid to pollen traces of sporadic taxa thought to have disappeared from a given area during the last glacial and post-glacial period.

The cultivation of Castanea sativa (Mill.) in Europe, from its origin to its diffusion on a continental scale

The history of Castanea sativa (sweet chestnut) cultivation since medieval times has been well described on the basis of the very rich documentation available. Far fewer attempts have been made to give a historical synthesis of the events that led to the cultivation of sweet chestnut in much earlier times. In this article we attempt to reconstruct this part of the European history of chestnut cultivation and its early diffusion by use of different sources of information, such as pollen studies, archaeology, history and literature. Using this multidisciplinary approach, we have tried to identify the roles of the Greek and Roman civilizations in the dissemination of chestnut cultivation on a European scale. In particular, we show that use of the chestnut for food was not the primary driving force behind the introduction of the tree into Europe by the Romans. Apart from the Insubrian Region in the north of the Italian peninsula, no other centre of chestnut cultivation existed in Europe during the Roman period. The Romans may have introduced the idea of systematically cultivating and using chestnut. In certain cases they introduced the species itself; however no evidence of systematic planting of chestnut exists. The greatest interest in the management of chestnut for fruit production most probably developed after the Roman period and can be associated with the socio-economic structures of medieval times. It was then that self-sufficient cultures based on the cultivation of chestnut as a source of subsistence were formed.