•A typology of nutritional functioning of forest ecosystems emerges from our study.•Forest productivity can be high even when pools of nutrients in soil are low.•Chemical fertility of forest ...ecosystems needs to encompass biogeochemical cycling.•Biological cycling becomes predominant when the soil nutrient reservoir is small.
Many forest ecosystems are developed on acidic and nutrient-poor soils and it is not yet clearly understood how forests sustain their growth with low nutrient resources. In forestry, the soil chemical fertility is commonly defined, following concepts inherited from agronomy, as the pool of plant-available nutrients in the soil at a given time compared to the nutritional requirement of the tree species. In this two-part study, Part 1 (Hansson et al., 2020) showed, through the compiled dataset of 49 forest ecosystems in France, Brazil and Republic of Congo, the limits of this definition of soil chemical fertility in forest ecosystem contexts. In this study (Part 2), we investigated the nutrient pools and fluxes between the different ecosystem compartments at 11 of the 49 sites in order to better characterize the role of the biogeochemical cycling of nutrients in the chemical fertility of forest ecosystems, and in particular the roles of the biological and geochemical components of biogeochemical cycling.
The analysis of our dataset shows different types of biogeochemical functioning. When the geochemical component (inputs through mineral weathering and/or atmospheric inputs, capillary rise) is predominant, sufficient nutrients are provided to the plant-soil system to ensure tree nutrition and growth. Conversely, when the geochemical component of the cycle brings too few nutrients to the plant-soil system, the biological component (litterfall, plant internal cycling) becomes predominant in tree nutrition and growth. In the latter case, forest production may be high even when pools of nutrients in the soil reservoir are low because small but active nutrient fluxes may continuously replenish the soil reservoir or may directly ensure tree nutrition by bypassing the soil reservoir.
This study highlights the necessity to include biogeochemical cycling and recycling fluxes in the definition and diagnosis methods of soil chemical fertility in forest ecosystems. We show that the chemical fertility is not only supported by the soil in forest ecosystem but by the sum of all the ecosystem’s compartments and fluxes between these pools.
Since the late 1970s, several long‐term ecological studies were conducted to better understand the biogeochemical functioning of Norway spruce stands in the Ardennes as these nutrient‐poor ecosystems ...were subject to high levels of acid deposition and exhibited symptoms of tree health decline. Between 1978 and 2009, acid deposition declined sharply, especially sulfur and to a lesser extent nitrogen deposition. The aim of this study was (i) to determine if the Norway spruce stands recovered after the reduction of acid deposition and (ii) to explain why such a recovery occurred or not. Therefore, we collected data from different projects carried out in the Ardennes to characterize the long‐term temporal trends in soil solution chemistry, foliar nutrition, and crown condition. In parallel, a model describing the nutrient cycling in forests (NuCM) was calibrated and used to check the consistency of the observed temporal trends and to explain them. The soil solution concentration of most of the elements decreased between 1978 and 2002, which was ascribed to a decrease in atmospheric deposition. For potassium, a decline in the exchangeable pool was also showed based on the simulation carried out with NuCM. As nitrogen (N) deposition remained at an elevated level, Norway spruce stands were progressively saturated in N and mineral nutrition became more and more unbalanced. Except the foliar N and Al concentration that remained constant and increased respectively, the foliar concentration of all other nutrients decreased between 1993 and 2009, which can be explained by the decrease in ion concentration in solution. These nutritional disorders weakened trees and were probably exacerbated during the 2003 summer drought, after which symptoms of vitality loss progressively appeared. In these N‐saturated ecosystems, the N cycle was disrupted by this health decline, which increased NO3− leaching reinforcing soil acidification and risk of aluminum (Al) toxicity.
As soil minerals are the principal input of nutrients in non-fertilized forests, the parameters which influence their dissolution must be determined to predict ecosystem sustainability. Notably, ...biological activities within the rhizosphere, such as root and micro-organism exudation and respiration, considerably affect mineral dissolution rate. Numerous laboratory studies have even demonstrated that certain biological processes involved in mineral weathering can be stimulated in low-nutrient availability conditions, resulting in an improvement of plant nutrition. The objective of this work was to determine in the field if the mineral dissolution rate linked to root and root-associated micro-organism activity is increased in low-nutrient availability conditions. Here, the impact of the rhizosphere on the dissolution of test minerals containing Ca (fluorapatite and labradorite plagioclase) was assessed in an acid forest soil in two stands of mature beeches (
Fagus sylvatica) presenting two levels of Ca availability: a control plot as well as a plot fertilized with Ca. Mineral-test bags were inserted at three different depths (−
2.5, −
10 and −
20 cm) in the control and the Ca-fertilized plots into both a zone with roots as well as a zone where roots had been excluded, thus permitting to assess the effect of the rhizosphere on the mineral dissolution. After four years of incubation in the soil, the minerals were weighed and observed by scanning electron microscope. In the control stand, linear dissolution voids were only observed on the mineral surfaces incubated in the zone with roots, suggesting that local biological activities occurring in the rhizosphere affect mineral weathering. This positive effect of the rhizosphere in the control stand was confirmed by quantification of the mineral dissolution, which revealed an increase of fluorapatite and labradorite weathering, reaching factors 3 to 4 at 20-cm depth. In contrast, the beech rhizosphere did not increase mineral dissolution, hyphae colonisation or linear dissolution marks in the Ca-fertilized stand. These results suggest that the rhizospheric biological activities acting on mineral weathering could be regulated by the nutrient availability in the ecosystem. This plasticity of the rhizospheric biological activities may thus contribute to the maintenance of ecosystem sustainability.
Reproduction in mammals is controlled by the hypothalamo-pituitary-gonadal (HPG) axis under the influence of external and internal factors such as photoperiod, stress, nutrition, and social ...interactions. Sheep are seasonal breeders and stop mating when day length is increasing (anestrus). However, interactions with a sexually active ram during this period can override the steroid negative feedback responsible for the anoestrus state, stimulate luteinizing hormone (LH) secretion and eventually reinstate cyclicity. This is known as the "ram effect" and research into the mechanisms underlying it is shedding new light on HPG axis regulation. The first step in the ram effect is increased LH pulsatile secretion in anestrus ewes exposed to a sexually active male or only to its fleece, the latter finding indicating a "pheromone-like" effect. Estradiol secretion increases in all ewes and this eventually induces a LH surge and ovulation, just as during the breeding season. An exception is a minority of ewes that exhibit a precocious LH surge (within 4 h) with no prior increase in estradiol. The main olfactory system and the cortical nucleus of the amygdala are critical brain structures in mediating the ram effect since it is blocked by their inactivation. Sexual experience is also important since activation (increased c-fos expression) in these and other regions is greatly reduced in sexually naïve ewes. In adult ewes kisspeptin neurons in both arcuate and preoptic regions and some preoptic GnRH neurons are activated 2 h after exposure to a ram. Exposure to rams also activates noradrenergic neurons in the locus coeruleus and A1 nucleus and increased noradrenalin release occurs in the posterior preoptic area. Pharmacological modulation of this system modifies LH secretion in response to the male or his odor. Together these results show that the ram effect can be a fruitful model to promote both a better understanding of the neural and hormonal regulation of the HPG axis in general and also the specific mechanisms by which male cues can overcome negative steroid feedback and trigger LH release and ovulatory cycles.
Sheep are seasonal breeders, but exposure of anoestrus ewes to rams results in a rapid increase in luteinizing hormone (LH) secretion, eventually leading to surge in LH. Although LH secretion is ...known to be under the control of many neurotransmitters, noradrenaline (NA) is of particular importance for the LH surge in induced ovulators, although little is known about its role in LH secretion induced by males in spontaneous ovulators. To address this question, anoestrus ewes fitted with guide-tubes in the medial preoptic area (MPOA) or the ventromedial hypothalamus were subjected to microdialysis and blood sampling every 15 min for an hour before and 2 h after exposure to rams, and the concentrations of LH, monoamine and amino acid transmitters were measured. In ewes implanted in the posterior MPOA that responded to the ram by an increase in LH pulses, NA concentrations changed after exposure to the ram (P<0.018) and were higher at 15 (P<0.054) and 45 min (P<0.03) after male introduction than before. By contrast, no change in NA could be detected in ewes implanted in the same region, but not responding to the ram, or in those showing increased LH pulsatility, but implanted in the anterior MPOA or in the ventromedial hypothalamus. No changes were observed in other neurotransmitters or when the ewes were exposed to male odour alone. These results suggest that NA release in the posterior MPOA is selectively involved in the triggering of LH secretion by rams in anoestrus ewes.
Abstract
Exposure to a ram during spring stimulates luteinizing hormone (LH) secretion and can induce ovulation in sexually quiescent ewes (“ram effect”). Kisspeptin (Kiss) present in the arcuate ...nucleus (ARC) and the preoptic area (POA) is a potent stimulators of LH secretion. Our aim was to investigate whether Kiss neurons mediate the increase in LH secretion during the ram effect. With double immunofluorescent detection, we identified Kiss neurons (Kiss IR) activated (Fos IR) by exposure to a ram for 2 hours (M2) or 12 hours (M12) or to ewes for 2 hours (C). The density of cells Kiss + Fos IR and the proportion of Kiss IR cells that were also Fos IR cells were higher in M2 and M12 than in C in ARC (P < 0.002) and POA (P < 0.02). In ARC, these parameters were also higher in M12 than in M2 (P < 0.02 and P < 0.05). Kiss antagonist (P234 10−6M) administered by retrodialysis in POA for 3 hours at the time of introduction of the ram reduced the amplitude of the male-induced increase in LH concentration compared with solvent (P < 0.02). In ARC, P234 had a more limited effect (P < 0.038 1 hour after P234) but pulse frequency increased less than after solvent (P = 0.07). In contrast, Kiss antagonist (P271 10−4M) infused in ARC but not POA 6 to 18 hours after introduction of the ram prevented the LH surge in the ewe (0/6 vs 4/5 and 4/6 in C). These results suggest that both populations of Kiss neurons are involved in the ram-induced pulsatile LH secretion and in the LH surge.
Both ARC and POA Kiss neurons are involved in the pulsatile LH secretion induced in anestrus ewes exposed to sexually active rams, but only ARC Kiss neurons are necessary for the ram-induced LH surge.
During spring sheep do not normally ovulate but exposure to a ram can induce ovulation. In some ewes an LH surge is induced immediately after exposure to a ram thus raising questions about the ...control of this precocious LH surge. Our first aim was to determine the plasma concentrations of oestradiol (E2) E2 in anoestrous ewes before and after the "ram effect" in ewes that had a "precocious" LH surge (starting within 6 hours), a "normal" surge (between 6 and 28h) and "late» surge (not detected by 56h). In another experiment we tested if a small increase in circulating E2 could induce an LH surge in anoestrus ewes. The concentration of E2 significantly was not different at the time of ram introduction among ewes with the three types of LH surge. "Precocious" LH surges were not preceded by a large increase in E2 unlike "normal" surges and small elevations of circulating E2 alone were unable to induce LH surges. These results show that the "precocious" LH surge was not the result of E2 positive feedback. Our second aim was to test if noradrenaline (NA) is involved in the LH response to the "ram effect". Using double labelling for Fos and tyrosine hydroxylase (TH) we showed that exposure of anoestrous ewes to a ram induced a higher density of cells positive for both in the A1 nucleus and the Locus Coeruleus complex compared to unstimulated controls. Finally, the administration by retrodialysis into the preoptic area, of NA increased the proportion of ewes with an LH response to ram odor whereas treatment with the α1 antagonist Prazosin decreased the LH pulse frequency and amplitude induced by a sexually active ram. Collectively these results suggest that in anoestrous ewes NA is involved in ram-induced LH secretion as observed in other induced ovulators.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
The proportion of anoestrous ewes ovulating after exposure to a sexually active ram is variable mainly due to whether an LH surge is induced. The aim of this study was to determine the role of ...oestradiol (E2) in the ram-induced LH surge. In one study, we measured the plasma concentrations of E2 in ewes of different breeds before and after the 'ram effect' and related these patterns to the presence and latency of the LH surge, while another compared ovarian responses with the 'ram effect' following exposure to rams for 2 or 12 h. In all ewes, the concentration of E2 increased 2-4 h after rams were introduced and remained elevated for 14.5 ± 0.86 h. The quantity of E2 secreted before the LH surge varied among breeds as did the mean concentration of E2. The granulosa cells of IF ewes collected after 12 h exposure to rams secreted more E2 and progesterone and had higher levels of StAR than the 2 h group but in MV ewes there was no differences between these groups for any of these parameters. These results demonstrate that the LH surge induced by the rams is a result of increased E2 secretion associated with increased levels of STAR in granulosa cells and that these responses varied among breeds. The results suggest that the variable occurrence of a LH surge and ovulation may be the result of variable ovarian responses to the 'ram effect' and insensitivity of the hypothalamus to the E2-positive feedback signal.
•A unique dataset combining soil properties and stand productivity was compiled.•Common soil chemical properties were poor predictors of stand productivity.•Forest soils with low nutrient stocks ...sometimes exhibit high stand productivity.•Nutrient fluxes and cycling could better explain stand productivity.
Forest soil fertility can be defined as a combination of physical, chemical and biological factors characterising the biomass production capacity of the soil. However, numerous ecological variables affect tree growth and the aim of the present study was to investigate the specific influence of soil chemical properties on tree productivity at 49 acidic forest sites. A standardized tree productivity index based on tree height expressed as dominant height of the studied stand divided by maximum tree height observed at the same age for the same species in the same climatic region was firstly computed at each site. This index is assumed to limit the influence of species, ages and climate. A soil database was also compiled with data on soil properties from 47 temperate (France) and two tropical (Congo, Brazil) sites. Data included seven tree species, varying in age from 1 to 175 years. Commonly used indicators such as C:N ratio, soil pH, as well as available and total pools of soil nutrients were compared to the standardized tree productivity index, to find the most reliable indicator(s). Nutrient pools at fixed mineral soil depths (down to 100 cm) were used, as well as (for 11 stands) the depth comprising 95% of fine roots.
Our results show that none of the common soil chemical parameters tested in this paper could individually explain stand productivity. Combinations of different parameters were also tested using PCA and they could better explain the variability of the data set but without being able to separate the sites according to their standardized tree productivity index. Moreover, random Forests performed on our dataset were unable to properly predict the standardized tree productivity index. Our results reinforce the idea that the influence of the soil chemical fertility on stand productivity is complex and the soil chemical parameters alone (individually or combined) are poor predictors of tree productivity as assessed by the H0:Hmax index. In this paper we focused on static soil chemical indicator and more dynamic indictors, such as nutrient fluxes involved in the biogeochemical cycles, could better explain stand productivity. A companion paper (Legout et al., 2020) focuses on the connection between productivity and different components of the biogeochemical cycle, using data from 11 of the stands presented in this paper.
► Brain structures activated by after contact with males or females in rams with different libidos or sexual preferences. ► Direct or indirect contact with a preferred sexual partner induced Fos in ...several diencephalic and cortical structures. ► Indirect contact with males did not induce Fos immunoreactivity in males interested in females. ► Activation in the preoptic area, the paraventricular nucleus and the medial BNST depended on the level of sexual activity. ► Preoptic area was more activated in males that displayed ejaculation than in those that displayed only courtship behaviour.
Using Fos, as a marker, we analysed the brain structures of rams, with different libidos or sexual preferences that had been activated by contact with males or females. Ile de France rams aged from 1.5 to 7 years were used. Fos immunoreactivity (Fos IR) was analysed in rams with high (HL) or low libido (LL) after 90
min of direct contact with females (HL DirF
n
=
7 or LL DirF
n
=
7) or in rams of high libido having indirect contact through a fence, with females (HL IndF
n
=
6) or males (HL IndM
n
=
5) and finally, in males who preferred other males as partners by indirect contact through a fence with males (MO IndM
n
=
4). Direct or indirect contact with a preferred sexual partner (LL DirF, HL Dir F, HL IndF, MO IndM) induced the appearance of Fos-IR cells in several diencephalic and cortical structures. Conversely, indirect contact with males did not induce Fos-IR in males interested in females (HL IndM). In the medial preoptic area (MPOA), the paraventricular nucleus and the medial bed nucleus of the stria terminalis the cell density of Fos IR cells was higher in HL Dir F than in LL DirF suggesting involvement in sexual motivation whereas only the MPOA seemed involved the consummatory component of sexual behaviour (Fos IR density HL DirF
>
HL IndF). The enthorinal cortex was the only structure specifically activated by males attracted to other males (Fos IR density MO IndM
>
HL IndM) whereas Fos IR density did not differ between the HL IndF and HL IndM groups.
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