N2-fixation and residual N effect of four legume species and four companion grass species

Inclusion of forage legumes in low-input forage mixtures improves herbage production and soil fertility through addition of nitrogen (N) from N2-fixation. The impact of different grass-legume mixtures on the N contribution of the forage mixture has rarely been investigated under comparable soil and climatic conditions. We conducted a field experiment on a sandy soil at two nitrogen levels with seven two-species forage mixtures: alfalfa, bird’s-foot trefoil, red clover, or white clover in mixture with perennial ryegrass, and white clover in mixture with meadow fescue, timothy, or hybrid ryegrass. We found high N2-fixation of more than 300 kg N ha-1 from both red clover and alfalfa even when the two mixtures received 300 kg total-N ha-1 in cattle slurry. The addition of cattle slurry N fertilizer lowered N2-fixation for white clover and red clover as expected, but for bird’s-foot trefoil and alfalfa no changes in the proportion of N derived from N2-fixation was observed. We conclude that the competition for available soil N from perennial ryegrass in mixture was an important factor for the proportion of N in alfalfa, white clover, and bird’s-foot trefoil obtained from N2-fixation. White clover had a high proportion of N derived from atmosphere for all companion grasses despite significant differences in white clover proportion. Although the perennial ryegrass-alfalfa mixture in the grass phase yielded more than twice the N from N2-fixation compared to white clover in the perennial ryegrass mixture, this did not in the following year lead to higher residual N effects of alfalfa. Both in terms of N yield in the grass phase and N yield in the subsequent spring barley red clover contributed most to the improvement of soil N fertility

The secret life of wild animals revealed by accelerometer data: how landscape diversity and seasonality influence the behavioural types of European hares

Context Landscape composition and configuration, as well as seasonal landscape dynamics shape the behaviour, movement and energy expenditure of animals, i.e. foraging, hiding or fleeing, and ultimately survival. Especially in highly modified agricultural systems, it is crucial to understand how animal behaviour is influenced by landscape context to develop sustainable land management concepts. Objectives We show how landscape composition and configuration, together with seasonal dynamics affect animal behavioural types, accounting for the different life-history events in both sexes. Methods We investigated 34 European hares in two contrasting agricultural landscapes (a simple and a complex landscape) by using tri-axial accelerometer data to classify the animals’ behaviour into five categories: resting, foraging, moving, grooming and standing upright (i.e. vigilance behaviour). We tested whether the amount of behaviours per category changed with landscape composition and configuration, season and sex. Results During peak breeding, hares in areas of high habitat diversity rested more, moved less and spent less time searching for resources. During winter, hares moved more and rested less. Females rested less and foraged more in areas with large agricultural fields. Conclusions A complex landscape is particularly important during the breeding season, allowing animals to allocate enough energy into reproduction. In winter, hares in areas of low habitat diversity may not find enough thermal and anti-predator shelter to move as much as they would need to meet their requirements. Hence, high habitat diversity and small field sizes guarantee species persistence in human-altered agricultural areas throughout the year

Circadian rhythms regulate the environmental responses of net CO2 exchange in bean and cotton canopies

Studies on the dependence of the rates of ecosystem gas exchange on environmental parameters often rely on the up-scaling of leaf-level response curves ('bottom-up' approach), and/or the down-scaling of ecosystem fluxes ('top-down' approach), where one takes advantage of the natural diurnal covariation between the parameter of interest and photosynthesis rates. Partly independent from environmental variation, molecular circadian clocks drive ∼24 h oscillations in leaf-level photosynthesis, stomatal conductance and other physiological processes in plants under controlled laboratory conditions. If present and of sufficient magnitude at ecosystem scales, circadian regulation could lead to different results when using the bottom-up approach (where circadian regulation exerts a negligible influence over fluxes because the environment is modified rapidly) relative to the top-down approach (where circadian regulation could affect fluxes as it requires the passage of a few hours). Here we dissected the drivers of diurnal net CO2 exchange in canopies of an annual herb (bean) and of a perennial shrub (cotton) through a set of experimental manipulations to test for the importance of circadian regulation of net canopy CO2 exchange, relative to that of temperature and vapor pressure deficit, and to understand whether circadian regulation could affect the derivation of environmental flux dependencies. Contrary to conventional wisdom, we observed how circadian regulation exerted controls over net CO2 exchange that were of similar magnitude to the controls exerted by direct physiological responses to temperature and vapor pressure deficit. Diurnal patterns of net CO2 exchange could only be explained by considering effects of environmental responses combined with circadian effects. Consequently, we observed significantly different results when inferring the dependence of photosynthesis over temperature and vapor pressure deficit when using the top-down and the bottom up approaches.We remain indebted to E. Gerardeau, D. Dessauw, J. Jean, P. Prudent (Aïda CIRAD), J.-J. Drevon, C. Pernot (Eco&Sol INRA), B. Buatois, A. Rocheteau (CEFE CNRS), A. Pra, A. Mokhtar and the full Ecotron team, in particular C. Escape, for outstanding technical assistance during experiment set-up, plant cultivation and measurements. Earlier versions of the manuscript benefitted from comments by M. Dietze, B. Medlyn, R. Duursma and Y.-S. Lin. This study benefited from the CNRS human and technical resources allocated to the ECOTRONS Research Infrastructures as well as from the state allocation ‘Investissement d'Avenir’ ANR-11-INBS-0001, ExpeER Transnational Access program, Ramón y Cajal fellowships (RYC-2012-10970 to VRD and RYC-2008-02050 to JPF), the Erasmus Mundus Master Course Mediterranean Forestry and Natural Resources Management (MEDfOR) and internal grants from UWS-HIE to VRD and ZALF to AG. We thank the Associate Editor T. Vesala and two anonymous reviewers for their help to improve this manuscript

Globalisation as a challenge or opportunity for organic farming

During one intensive week in October 2005, the authors were gathered to discuss the impact that globalisation has on the Organic Food Systems and the opportunities that globalisation opens up for developing these systems. The meeting took place as a Ph.D. course under the auspices of the Research School of Organic Farming and Food Systems (SOAR; www.soar.dk). All participants research within Organic Agriculture and Food Production in one way or another

Night and day - Circadian regulation of night-time dark respiration and light-enhanced dark respiration in plant leaves and canopies

The potential of the vegetation to sequester C is determined by the balance between assimilation and respiration. Respiration is under environmental and substrate-driven control, but the circadian clock might also contribute. To assess circadian control on night-time dark respiration (RD) and on light enhanced dark respiration (LEDR) - the latter providing information on the metabolic reorganization in the leaf during light-dark transitions - we performed experiments in macrocosms hosting canopies of bean and cotton. Under constant darkness (plus constant air temperature and air humidity), we tested whether circadian regulation of RD scaled from leaf to canopy respiration. Under constant light (plus constant air temperature and air humidity), we assessed the potential for leaf-level circadian regulation of LEDR. There was a clear circadian oscillation of leaf-level RD in both species and circadian patterns scaled to the canopy. LEDR was under circadian control in cotton, but not in bean indicating species-specific controls. The circadian rhythm of LEDR in cotton might indicate variable suppression of the normal cyclic function of the tricarboxylic-acid-cycle in the light. Since circadian regulation is assumed to act as an adaptive memory to adjust plant metabolism based on environmental conditions from previous days, circadian control of RD may help to explain temporal variability of ecosystem respiration.This study benefited from the CNRS human and technical resources allocated to the ECOTRONS Research Infrastructures as well as from the state allocation ‘Investissement d'Avenir’ AnaEE-France ANR-11-INBS-0001, ExpeER Transnational Access program, Ramón y Cajal fellowships (RYC-2012-10970 to VRD and RYC-2008-02050 to JPF), the Erasmus Mundus Master Course MEDfOR, internal grants from UWS-HIE to VRD and ZALF to AG and Juan de la Cierva-fellowships (IJCI-2014-21393 to JGA). We remain indebted to E. Gerardeau, D. Dessauw, J. Jean, P. Prudent (Aïda CIRAD), J.-J. Drevon, C. Pernot (Eco&Sol INRA), B. Buatois, A. Rocheteau (CEFE CNRS), A. Pra, A. Mokhtar and the full Ecotron team, in particular C. Escape, for outstanding technical assistance

Nitrogen transfer from forage legumes to nine neighbouring plants in a multi-species grassland

Legumes play a crucial role in nitrogen supply to grass-legume mixtures for ruminant fodder. To quantify N transfer from legumes to neighbouring plants in multi-species grasslands we established a grass-legume-herb mixture on a loamy-sandy site in Denmark. White clover (Trifolium repens L.), red clover (Trifolium pratense L.) and lucerne (Medicago sativa L.) were leaf-labelled with 15N enriched urea during one growing season. N transfer to grasses (Lolium perenne L. and xfestulolium), white clover, red clover, lucerne, birdsfoot trefoil (Lotus corniculatus L.), chicory (Cichorium intybus L.), plantain (Plantago lanceolata L.), salad burnet (Sanguisorba minor L.)and caraway (Carum carvi L.) was assessed. Neighbouring plants contained greater amounts of N derived from white clover (4.8 gm-2) compared with red clover (2.2 gm-2) and lucerne (1.1 gm-2). Grasses having fibrous roots received greater amounts of N from legumes than dicotyledonous plants which generally have taproots. Slurry application mainly increased N transfer from legumes to grasses. During the growing season the three legumes transferred approximately 40 kg N ha-1 to neighbouring plants. Below-ground N transfer from legumes to neighbouring plants differed among nitrogen donors and nitrogen receivers and may depend on root characteristics and regrowth strategies of plant species in the multi-species grassland

Performance of multi-species grassland mixtures. Mineral accumulation, nitrogen transfer and nitrogen uptake

This thesis is located within the framework of research on temperate agricultural grasslands. Plant diversity in temperate agricultural grasslands is impoverished today. Many conventional grasslands in Europe consist of simple grass mixtures, sometimes monocultures. Organic and increasingly conventional grasslands are sown as mixtures of grasses and legumes and from time to time as multi-species mixtures that include forage herbs. Temporary grasslands are regularly resown or incorporated in a crop rotation and produce ruminant fodder. Some studies indicate that various forage herbs have traits that may complement grass-legume mixtures; among others, increased macro- and micromineral concentration. Hence multi-species grassland mixtures, comprising grasses, legumes and herbs, may have a potential to improve common grasslands consisting of low plant diversity. The legume component of a grass-legume mixture contributes to nitrogen (N) self-sufficiency of the grassland because of its ability for symbiotic N2 fixation, which allows reduced fertilizer application. Several studies present a positive plant diversityproductivity relationship for temporary European grasslands. Increased soil nutrient use in grasslands with higher plant diversity may be one explanation, where, for example, deep-rooted plant species enable nutrient uptake from deep soil layers. In a state of limbo between ecological hypotheses and agricultural needs, and with the background of limited information on forage herbs and multi-species grassland mixtures, the objective of this Ph.D. study was to quantify the performance of multi-species grassland mixtures in terms of three nutrient processes: 1) mineral accumulation, 2) nitrogen transfer and 3) nitrogen uptake. Three field experiments were conducted in Denmark: two on a loamy-sandy site and one on a sandy-loamy site. The four forage herbs – chicory (Cichorium intybus L.), plantain (Plantago lanceolata L.), salad burnet (Sanguisorba minor L.) and caraway (Carum carvi L.) – grown in a nine-species grass-legume-herb mixture showed in my first field experiment, potential in increasing the macro- and micromineral quality of ruminant fodder derived from grasslands. In a second experiment, legumes, which were confirmed to be the N-richest forage plants of the multi-species mixture, donated N to all nine neighboring plants. Grasses (xFestulolium and Lolium perenne L.) received the largest amount of N among the receiver plants, mainly from white clover (Trifolium repens L.), and slurry application mainly stimulated N transfer from legumes towards grasses. The third experiment confirmed the strong N absorption characteristic of perennial ryegrass when grown in a multispecies mixture. When deep-rooted chicory and lucerne (Medicago sativa L.) were added to a shallow-rooted perennial ryegrass-white clover mixture, total N uptake from three vertical soil layers of this four-species grass-legume-herb mixture did not increase. Chicory, however, took up greater proportions of N from deeper soil layers compared with the other plant species. This study showed that multi-species grassland mixtures have the potential to improve the common simple grass or grass-legume mixtures by reducing the need for external mineral supplementation and by more efficient N use. Yet to use varying plant diversity as a management tool in an agricultural system based on grasslands, more knowledge from grass-legume-herb mixtures to animal performance is needed. Ecologists, agronomists and seed companies may work together to develop improved sustainable grasslands which to a certain extent may mimic natural grasslands

Næringsstofdynamik i græsmarker med kløver og urter

Der er ikke megen viden om urter i omdriftsgræsmarker. For at undersøge næringsstofdynamikken i græsmarker med kløver og urter udførte jeg markforsøøg på forsøgsgården i Foulum i 2007 og 2008. Forsøgene viser at urter har et potentiale til at forbedre en kløvergræsmark

Adaptations for vision in dim light: impulse responses and bumps in nocturnal spider photoreceptor cells (Cupiennius salei Keys)

The photoreceptor cells of the nocturnal spider Cupiennius salei were investigated by intracellular electrophysiology. (1) The responses of photoreceptor cells of posterior median (PM) and anterior median (AM) eyes to short (2 ms) light pulses showed long integration times in the dark-adapted and shorter integration times in the light-adapted state. (2) At very low light intensities, the photoreceptors responded to single photons with discrete potentials, called bumps, of high amplitude (2-20 mV). When measured in profoundly dark-adapted photoreceptor cells of the PM eyes these bumps showed an integration time of 128 +/- 35 ms (n = 7) whereas in dark-adapted photoreceptor cells of AM eyes the integration time was 84 +/- 13 ms (n = 8), indicating that the AM eyes are intrinsically faster than the PM eyes. (3) Long integration times, which improve visual reliability in dim light, and large responses to single photons in the dark-adapted state, contribute to a high visual sensitivity in Cupiennius at night. This conclusion is underlined by a calculation of sensitivity that accounts for both anatomical and physiological characteristics of the eye