Segment-based coancestry,additive relationship and genetic variance within and between the Norwegian and the Swedish Fjord horse populations

ABSTRACT

The Fjord horse originates from Norway but forms a global population due to several small populations in foreign countries. There exists no information about the additive relationship and the genetic variance between these subpopulations. By collecting blood samples from Norwegian and Swedish Fjord horses, a sample of 311 Norwegian and 102 Swedish horses gave 485,918 SNPs available for analysis. Their inbreeding coefficients were calculated and compared to the pairwise coancestry and the shared genomic segments. The effective population size was almost similar with the two methods in the Norwegian Fjord horse population (63 and 71), but very different in the Swedish population (269 and 1136) and unprecise due to a much smaller number of observations. The study showed that coancestry from shared genomic segments can be used to estimate additive genetic relationship and genetic variation within and between the global populations of the Fjord horse.     

Assessment and up-scaling of CO2 exchange by patches of the herbaceous vegetation mosaic in a Portuguese cork oak woodland

Long-term eddy covariance measurements over a montado oak woodland in southern Portugal have documented a vulnerability to predicted decreases in springtime rainfall, since water availability during spring limits annual CO₂ gain, the growth of fodder for animals, and the production of cork by Quercus suber. The current study examined CO₂ exchange of three different herbaceous vegetation components distributed over montado landscapes and within the footprint of long-term landscape eddy covariance monitoring studies. Simultaneous measurements with eddy covariance at two sites and with manually operated chambers at multiple locations revealed that slow drainage of shallow basins, the onset of drying at higher sites and a high release of CO₂ below tree canopies significantly influenced the overall course of montado ecosystem gas exchange during the spring.Hyperbolic light response models were employed to up-scale and compare herbaceous gas exchange with landscape net ecosystem CO₂ flux. The up-scaling demonstrates the importance of the herbaceous understory in determining annual carbon balance of the montado and suggests a relatively small additional CO₂ uptake by the tree canopies and boles, i.e., by the aboveground tree compartment, during springtime. Annual flux totals obtained during the extremely dry year 2005 and a normal precipitation year 2006 for the oak woodland and a nearby grassland were essentially the same, indicating that both ecosystems similarly exploit available resources. Based on comparisons with additional temperate grasslands, we can visualize the montado herbaceous cover as a typical European grassland canopy, but where temperature fluctuations in winter control uptake, and where total production depends on springtime rainfall as it controls phenological events and eventually dieback of the vegetation. On the other hand, tree canopies remain active longer during late spring and early summer, modifying the montado response from that of grassland. Uncertainties in flux estimates via both chamber and eddy covariance methodologies currently prevent a full understanding of vegetation/atmosphere coupling, of the recycling of CO₂ between the understory communities and trees, and of relationships between exchange rates of individual components of the vegetation mosaic and overall carbon and water balances in montado landscapes.     

An analysis of light effects on foliar morphology,physiology, and light interception in temperate deciduous woody species of contrasting shade tolerance

Maximum Rubisco activities (V(cmax)), rates of photosynthetic electron transport (J(max)), and leaf nitrogen and chlorophyll concentrations were studied along a light gradient in the canopies of four temperate deciduous species differing in shade tolerance according to the ranking: Populus tremula L. < Fraxinus excelsior L. < Tilia cordata Mill. = Corylus avellana L. Long-term light environment at the canopy sampling locations was characterized by the fractional penetration of irradiance in the photosynthetically active spectral region (I(sum)). We used a process-based model to distinguish among photosynthesis limitations resulting from variability in fractional nitrogen investments in Rubisco (P(R)), bioenergetics (P(B), N in rate-limiting proteins of photosynthetic electron transport) and light harvesting machinery (P(L), N in chlorophyll and thylakoid chlorophyll-protein complexes). On an area basis, V(cmax) and J(max) (V(a) (cmax) and J(a) (max)) increased with increasing growth irradiance in all species, and the span of variation within species ranged from two (T. cordata) to ten times (C. avellana). Examination of mass-based V(cmax) and J(max) (V(m) (cmax) and J(m) (max)) demonstrated that the positive relationships between area-based quantities and relative irradiance mostly resulted from the scaling of leaf dry mass per area (M(A)) with irradiance. Although V(m) (cmax) and J(m) (max) were positively related to growth irradiance in C. avellana, and J(m) (max) was positively related to irradiance in P. tremula, the variation range was only a factor of two. Moreover, V(m) (cmax) and J(m) (max) were negatively correlated with relative irradiance in T. cordata. Rubisco activity in crude leaf extracts generally paralleled the gas-exchange data, but it was independent of light in T. cordata, suggesting that declining V(m) (cmax) with increasing relative irradiance was related to increasing diffusive resistances from the intercellular air spaces to the sites of carboxylation in this species. Because irradiance had little effect on foliar nitrogen concentration, the relationships of P(B) and P(R) with irradiance were similar to those of V(m) (cmax) and J(m) (max). Shade-intolerant species tended to have greater P(B) and P(R) and also larger V(a) (cmax) and J(a) (max) than more shade-tolerant species. However, for the whole material, P(B) and P(R) varied only about 50%, whereas V(a) (cmax) and J(a) (max) varied more than 15-fold, further emphasizing the importance of leaf anatomical plasticity in determining photosynthetic acclimation to high irradiance. Leaf chlorophyll concentrations and fractional nitrogen investments in light harvesting increased hyperbolically with decreasing irradiance to improve quantum use efficiency for incident irradiance. The effect of irradiance on P(L) was of the same order as its effect in the opposite direction on M(A), leading to either a constant model estimate of leaf absorptance with I(sum) or a slightly positive correlation. We conclude that leaf morphological plasticity is a more relevant determinant of foliage adaptation to high irradiance than foliage biochemical properties, whereas biochemical adaptation to low irradiance is of the same magnitude as the anatomical adjustments. Although shade-tolerant species did not have greater chlorophyll concentrations and P(L) than shade-intolerant species, they possessed lower M(A), and could maintain a more extensive foliar display for light capture with constant biomass investment in leaves.     

Annual and seasonal variations in photosynthetic capacity of Fagus crenata along an elevation gradient in the Naeba Mountains, Japan

Canopy photosynthetic capacity, measured as leaf maximum carboxylation rate (V (cmax)), is a key factor in ecosystem gas exchange models applied at different scales. We report seasonal and interannual variations in V(cmax) of natural beech stands (Fagus crenata Blume) along an altitudinal gradient in the temperate climate zone of Japan. Estimates are based on 6 years of gas exchange measurements. Pronounced seasonal and interannual variations in V(cmax) normalized to 25 degrees C (V(c,25)) were found for sun leaves. The seasonal pattern of V(c,25) generally followed an inverse parabolic curve, with an increase in spring, peak values in the middle of the growth period and a decline in autumn. Leaf nitrogen concentration (N(l)) and leaf mass per area were significantly related to V(c,25) during spring and summer, but were unrelated in autumn when V(c,25) declined. Annual peak V(c,25) ranged from 40.1 to 97.0 micromol m(-2) s(-1) and varied over as much as a twofold range at a particular site. Annual peak V(c,25) occurred about 28 days before annual peak N(l), with which it was poorly related. Our results show that it can be inappropriate to include constant values of photosynthetic parameters in ecosystem gas exchange models.     

A single drought event of 100‐year recurrence enhances subsequent carbon uptake and changes carbon allocation in experimental grassland communities

Evidence suggests that the expected increase in frequency and magnitude of extreme weather events during climate change will alter plant productivity. Therefore, extreme weather events might also be capable of changing C sequestration and allocation. Here, experimental grassland communities of two species compositions, differing in their diversity, were exposed either to a simulated single drought or to a heavy‐rainfall event. The magnitude of these manipulations imitated the local 100‐year weather extreme according to extreme‐value statistics. Effects on Net Ecosystem CO₂ Exchange (NEE in µmol m^–2 s^–1) as well as aboveground biomass production and leaf‐area index (LAI) were recorded from prior to the manipulations until two months after the manipulations ended. Initial light utilization efficiency and maximum NEE increased after the drought. No change in the respiration was detected and maximum uptake capacity (GPP_max) was 15% higher for the drought‐manipulated plots compared to controls, which indicates an enhanced CO₂ uptake into the systems. The level of diversity was also found to alter the light‐response curves, increasing respiration and maximum NEE to a higher degree than drought in the more diverse compared to the less diverse community. This resulted in an increase of GPP_max by 55%. No significant interactions between species composition and weather manipulations were detected. Interestingly, aboveground biomass production was not significantly affected by weather manipulations, even though LAI increased due to drought. This increase was caused by a decrease in the ratio between reproductive and vegetative growth. The heavy‐rainfall manipulation resulted in no significant effects. Our data suggest that C sequestration can be enhanced by a single weather event. However the importance, long‐term duration, and thresholds or turning points of such effects need to be investigated further as intensification of weather extremes is currently emerging as one of the most important facets of climate change.     

Physiological and morphological responses to water stress in two Acacia species from contrasting habitats

Container-grown seedlings of Acacia tortilis Forsk. Hayne and A. xanthophloea Benth. were watered either every other day (well watered) or every 7 days (water-stressed) for 1 year in a greenhouse. Total plant dry mass (T(dm)), carbon allocation and water relations were measured monthly. Differences in leaf area (LA) accounted for differences in T(dm) between the species, and between well-watered and water-stressed plants. Reduction in LA as a result of water stress was attributed to reduced leaf initiation, leaf growth rate and leaf size. When subjected to prolonged water stress, Acacia xanthophloea wilted more rapidly than A. tortilis and, unlike A. tortilis, lost both leaves and branches. These differences between species were attributed to differences in the allocation of carbon between leaves and roots and in the ability to adjust osmotically. Rapid recovery in A. xanthophloea following the prolonged water-stress treatment was attributed to high cell wall elasticity. Previous exposure to water stress contributed to water-stress resistance and improved recovery after stress.     

Age-related effects on leaf area/sapwood area relationships,canopy transpiration and carbon gain of Norway spruce stands (Picea abies) in the Fichtelgebirge,Germany

Stand age is an important structural determinant of canopy transpiration (E(c)) and carbon gain. Another more functional parameter of forest structure is the leaf area/sapwood area relationship, A(L)/A(S), which changes with site conditions and has been used to estimate leaf area index of forest canopies. The interpretation of age-related changes in A(L)/A(S) and the question of how A(L)/A(S) is related to forest functions are of current interest because they may help to explain forest canopy fluxes and growth. We conducted studies in mature stands of Picea abies (L.) Karst. varying in age from 40 to 140 years, in tree density from 1680 to 320 trees ha(-1), and in tree height from 15 to 30 m. Structural parameters were measured by biomass harvests of individual trees and stand biometry. We estimated E(c) from scaled-up xylem sap flux of trees, and canopy-level fluxes were predicted by a three-dimensional microclimate and gas exchange model (STANDFLUX). In contrast to pine species, A(L)/A(S) of P. abies increased with stand age from 0.26 to 0.48 m(2) cm(-2). Agreement between E(c) derived from scaled-up sap flux and modeled canopy transpiration was obtained with the same parameterization of needle physiology independent of stand age. Reduced light interception per leaf area and, as a consequence, reductions in net canopy photosynthesis (A(c)), canopy conductance (g(c)) and E(c) were predicted by the model in the older stands. Seasonal water-use efficiency (WUE = A(c)/E(c)), derived from scaled-up sap flux and stem growth as well as from model simulation, declined with increasing A(L)/A(S) and stand age. Based on the different behavior of age-related A(L)/A(S) in Norway spruce stands compared with other tree species, we conclude that WUE rather than A(L)/A(S) could represent a common age-related property of all species. We also conclude that, in addition to hydraulic limitations reducing carbon gain in old stands, a functional change in A(L)/A(S) that is related to reduced light interception per leaf area provides another potential explanation for reduced carbon gain in old stands of P. abies, even when hydraulic constraints increase in response to changes in canopy architecture and aging.     

Responses of foliar photosynthetic electron transport, pigment stoichiometry, and stomatal conductance to interacting environmental factors in a mixed species forest canopy

We studied limitations caused by variations in leaf temperature and soil water availability on photosynthetic electron transport rates calculated from foliar chlorophyll fluorescence analysis (U) in a natural deciduous forest canopy composed of shade-intolerant Populus tremula L. and shade-tolerant Tilia cordata Mill. In both species, there was a positive linear relationship between light-saturated U (Umax) per unit leaf area and mean seasonal integrated daily quantum flux density (Ss, mol per square m per day). Acclimation of leaf dry mass per area and nitrogen per area to growth irradiance largely accounted for this positive scaling. However, the slopes of the Umax versus Ss relationships were greater on days when leaf temperature was high than on days when leaf temperature was low. Overall, Umax varied 2.5-fold across a temperature range of 20-30 degrees C. Maximum stomatal conductance (Gmax) also scaled positively with Ss. Although Gmax observed during daily time courses, and stomatal conductances during Umax measurements declined in response to seasonally decreasing soil water contents, was insensitive to prolonged water stress, and was not strongly correlated with stomatal conductances during its estimation. These results suggest that photorespiration was an important electron sink when intercellular CO2 concentration was low because of closed stomata. Given that xanthophyll cycle pool size (VAZ, sum of violaxanthin, antheraxanthin, and zeaxanthin) may play an important role in dissipation of excess excitation energy, the response of VAZ to fluctuating light and temperature provided another possible explanation for the stable Umax. Xanthophyll cycle carotenoids per total leaf chlorophyll (VAZ/Chl) scaled positively with integrated light and negatively with daily minimum air temperature, whereas the correlation between VAZ/Chl and irradiance was best with integrated light averaged over 3 days preceding foliar sampling. We conclude that the potential capacity for electron transport is determined by long-term acclimation of U to certain canopy light conditions, and that the rapid adjustment of the capacity for excitation energy dissipation plays a significant part in the stabilization of this potential capacity. Sustained high capacity of photosynthetic electron transport during stress periods provides an explanation for the instantaneous response of U to short-term weather fluctuations, but also indicates that U restricts potential carbon gain under conditions of water limitation less than does stomatal conductance.     

Site-specific water relations and stomatal response of Quercus ilex in a Mediterranean watershed

Intraspecific variations in the water relations and stomatal response of Quercus ilex L. were analyzed under field conditions by comparing trees at two locations within a Mediterranean watershed (l'Avic, Catalonia, NE Spain). Distinct environmental gradients exist between the two sites (referred to as ridge top at 975 m and valley bottom at 700 m) with greater soil depth for water storage, reduced radiation, reduced wind and higher water vapor pressure deficits at the valley bottom than at the ridge top. Osmotic adjustment and changes in tissue elasticiity did not significantly increase drought resistance in the trees studied. The leaf-to-air vapor pressure difference (Deltaw) threshold for inducing stomatal closure was higher at the ridge top (15.6 kPa MPa(-1) +/- 0.5 SE) than at the valley bottom (9.8 kPa MPa(-1) +/- 1.0 SE). However, increases in Deltaw beyond the threshold were followed by greater reductions in leaf conductance of trees at the ridge top than at the valley bottom. At both sites, maximum leaf conductance was related to predawn shoot water potential which, in turn, was related to watershed stream flow. The effects of water deficits during the dry summer of 1989 were more severe in trees at the valley bottom than at the ridge top. During periods of high evaporative demand, site-specific differences in the control of water loss led to more conservative water use by trees at the ridge top and, thus, to even greater drought avoidance (higher predawn water potentials) in late summer.