Effects of increased temperature and nitrogen availability on the displacement of Calluna vulgaris by Pteridium aquilinum

Abstract. Atmospheric greenhouse gas concentrations will continue to rise for the foreseeable future, and, due to radiative forcing, surface temperatures will also increase. If soils warm sufficiently, this will result in increased litter decomposition rates, while nitrogen availability will also rise due to increased deposition. A field-based experiment manipulating air temperatures using polythene tents and nitrogen availability by adding 50 kg N ha^-1 yr^-1 has shown that this will result in further encroachment of Pteridium into Calluna moorland. Although this experiment, treating plots on and around the boundary between Pteridium and Calluna, did not show any statistically significant effect on the movement of the boundary, there were clear indications of Calluna dieback at the boundary. In addition, the frequency of Pteridium fronds increased in the Calluna-dominated areas, while Calluna became less abundant in the pure Pteridium plots. Calluna dieback was not due to litter accumulation as is frequently assumed, but the dieback is likely to be caused by shading of Calluna by the increased growth of Pteridium fronds. Botanical diversity was higher in the Pteridium plots, but considering the importance of grouse shooting for the maintenance of Calluna moorland, displacement of Calluna by Pteridium may result in a lower conservation and economic value of moorlands.     

Effects of cyanogenesis in bracken fern (Pteridium aquilinum) on associated insects

Abstract. 1. The relationship between cyanogenesis in bracken fern and the insect fauna feeding on the plant was investigated over a 3 year period. The most common insects between May and July, while cyanide levels were high, were the sawflies Strongylogaster impressata Provancher, S.multicincta Norton, Aneug-menus fzavipes (Norton), the aphid Macrosiphum euphorbiae (Thomas) and a microlepidopteran species of Monochroa .
2. Collections of insects from cyanogenic and acyanogenic fronds showed significantly fewer sawflies on the cyanogenic fronds. The aphid and the microlepidopteran were randomly distributed with respect to cyanogenicity.
3. Feeding tests for two of the sawfly species ( Simpressata and Smulticincta ) showed that larvae grew more slowly and had a higher mortality when raised on cyanogenic fronds than on acyanogenic ones.
4. Field collected cyanogenic bracken fronds were found to have sustained less damage from chewing herbivores compared with acyanogenic fronds.     

The morphology of bracken (Pteridium aquilinum (L.) Kuhn) in the North York Moors—a comparison of the mature stand and the interface with heather (Calluna vulgaris (L.) Hull) 1. The fronds

The encroachment of bracken (Pteridium aquilinum (L.) Kuhn) into areas previously dominated by heather represents a threat to the ecology, agricultural economy and landscape value of many UK upland areas, including the moorland of the North York Moors National Park. The morphology of bracken, within a mature stand and at several bracken-heather interfaces, has been studied at a number of sites within the National Park. Differences have been found in the frond growth of bracken in a mature stand, at stationary stand margins, and at advancing stand margins where bracken is encroaching into heather. Frequency of fronds present on bracken rhizome growing at a stationary stand margin close to the interface with heather (1–2 m behind the boundary) are approximately the same as those found within a mature stand. At advancing margins (again 1–2 m behind the boundary), maximum frond densities were often found to exceed those present in either a mature stand or at a stationary margin. Frond numbers decline rapidly at the stand margins as distance from the stand increases. This is especially true where the front is stationary and bracken is not encroaching into heather at a significant rate. Maximum frond heights in a mature stand consistently exceed those at stand margins (even 1–2 m into the stand) and are greater at stationary margins than at advancing margins. Outlying fronds at the edges of bracken stands are generally present in greater numbers, and further into the area dominated by heather, where the margin is advancing. Heights of outlying fronds fall as distance from the bracken stand increases, as does stipe length. Fronds at the edges of bracken stands emerge each spring before those further into the stand and are therefore particularly vulnerable to frost damage. Outlying fronds are not, however, the first to emerge. Early emerging fronds reach their maximum height and eventually become senescent before later emerging fronds. Whilst most fronds emerge before the end of June a few fronds continue to emerge throughout the summer. Frond densities close to the edges of bracken stands (1–2 m into the stand) are comparable to those in a mature stand. At advancing stand margins frond densities generally exceed those in a mature stand, suggesting that a large number of potential entry points for foliage-applied herbicides are available for bracken control at the stand margins. The ratio of potential uptake points to biomass of rhizome is also greatest at the edges of the stand, and the canopy 1–2 m into the stand is usually almost completely closed. It is possible therefore, the efficacy of herbicides could be improved by the use of small scale applications, using tractors or hand-held sprayers, close to the margins of bracken stands.     

The morphology of bracken (Pteridium aquilinum (L.) Kuhn) in the North York Moors—a comparison of the mature stand and the interface with heather (Calluna vulgaris (L.) Hull) 2. The rhizome

The rhizome system of mature bracken (Pteridium aquilinum (L.) Kuhn) contains large reserves of both biomass (mean 8.63 kg m?² fr. wt) and buds (mean 565 m^-2) which are largely responsible for both its persistence and its often rapid rates of vegetative encroachment. Within areas such as the North York Moors the spread of bracken into areas previously dominated by heather and grass is considered undesirable because of reduced land value in terms of both agriculture and ecological diversity. In this paper we describe the morphology of bracken rhizome within a mature bracken stand, and at advancing and stationary stand margins where bracken-heather interfaces occur. Stationary margins, i.e. those where bracken is not encroaching into heather at a significant rate, often have morphological characteristics intermediate to those of a mature stand and an advancing margin. In the mature stand rhizome biomass is dominated by carbohydrate-storing long shoots which comprise 63% of the total fresh weight, whilst the majority of rhizome buds (89% of all active and 86% of all dormant buds) are found on frond-bearing short shoots. At the margins of a bracken stand the proportion of rhizome which is composed of long shoots is even greater, and that of short shoots small relative to that in the mature stand. More transitional shoots are also found at the stand margins. Hence close to the margin a greater proportion of fronds is found on transitional rhizome than is the case in more mature parts of the stand. The majority of buds on all types of rhizome are in a dormant state. The proportion of buds which are active is, however, greater on long and transitional shoots than on short shoots. Hence, a larger proportion of buds are active close to the margin where the rhizome is composed less of short shoots than is the case further into a mature stand. The differences in the morphology of bracken in a mature stand and at the stand margins which are identified here support the idea of controlling bracken at stand margins in preference to the spraying of large areas of dense, mature bracken. Morphological differences include an increased proportion of active buds, greater frequency of fronds per unit rhizome biomass, reduced biomass reserves. Improved conditions for the re-invasion and re-establishment of alternative vegetation are also available at stand margins in comparison with the centre of a dense bracken stand.     

Selection on leaf ecophysiological traits in a desert hybrid Helianthus species and early-generation hybrids

Leaf ecophysiological traits related to carbon gain and resource use are expected to be under strong selection in desert annuals. We used comparative and phenotypic selection approaches to investigate the importance of leaf ecophysiological traits for Helianthus anomalus, a diploid annual sunflower species of hybrid origin that is endemic to active desert dunes. Comparisons were made within and among five genotypic classes: H. anomalus, its ancestral parent species (H. annuus and H. petiolaris), and two backcrossed populations of the parental species (designated BC2ann and BC2pet) representing putative ancestors of H. anomalus. Seedlings were transplanted into H. anomalus habitat at Little Sahara Dunes, Utah, and followed through a summer growing season for leaf ecophysiological traits, phenology, and fitness estimated as vegetative biomass. Helianthus anomalus had a unique combination of traits when compared to its ancestral parent species, suggesting that lower leaf nitrogen and greater leaf succulence might be adaptive. However, selection on leaf traits in H. anomalus favored larger leaf area and greater nitrogen, which was not consistent with the extreme traits of H. anomalus relative to its ancestral parents. Also contrary to expectation, current selection on the leaf traits in the backcross populations was not consistently similar to, or resulting in evolution toward, the current H. anomalus phenotype. Only the selection for greater leaf succulence in BC2ann and greater water-use efficiency in BC2pet would result in evolution toward the current H. anomalus phenotype. It was surprising that the action of phenotypic selection depended greatly on the genotypic class for these closely related sunflower hybrids grown in a common environment. We speculate that this may be due to either phenotypic correlations between measured and unmeasured but functionally related traits or due to the three genotypic classes experiencing the environment differently as a result of their differing morphology.     

Effects of long-term exposure to elevated CO2 and increased nutrient supply on bracken (Pteridium aquilinum)

1. Bracken ( Pteridium aquilinum ) is an important fern with a global distribution. Little is known of the response of this species to elevated CO₂. We investigated the effects of high CO₂ (570 compared with 370 μmol mol^–1) with and without an increased nutrient supply (a combined N, P, K application) on the growth and physiology of bracken, growing in containers in controlled-environment glasshouses, over two full growing seasons. Results of growth and physiology determinations are reported for the second season.
2. Elevated CO₂ had little impact on the growth or allocation of dry mass in bracken. No significant changes were detected in dry mass of the total plant or any of the organs: rhizomes, roots and fronds. In contrast to the small effects of high CO₂, the high nutrient treatment caused a three-fold stimulation of total plant dry mass and an increase in the allocation of dry mass to above ground when compared with low nutrient controls.
3. Net photosynthetic rates in saturating light were increased by both high CO₂ and nutrient treatments, particularly in spring months (May and June). Growth in elevated CO₂ did not cause a down-regulation in light-saturated rates of photosynthesis. The increased carbon gain in the high CO₂ treatments was accompanied, in the low-nutrient plants, by higher concentrations of carbohydrates. However, in high-nutrient plants the CO₂ treatment did not cause an accumulation of carbohydrates. The absence of a growth response to elevated CO₂ in bracken despite significant increases in photosynthesis requires further investigation.     

Estimates of gene flow, genetic substructure and population heterogeneity in bracken (Pteridium aquilinum)

Bracken [ Pteridium aquilinum (L.) Kuhn] is a cosmopolitan species and is a noxious weed in many areas. Because of its abundance, particularly in Britain, bracken affords an ideal system for investigating various aspects of population genetics and evolution. High mobility of dispersal units (spores) suggests that rates of gene flow among distant populations should be high. Gene flow is a major evolutionary force that influences the genetic structure of populations. To examine the effects of gene flow on population heterogeneity and population substructuring in bracken, starch gel electrophoresis of enzymes was used to provide the necessary genetic database. Allele frequency data at 21 loci were obtained for seven British populations, one Majorcan and one from the eastern United States. A model was employed to estimate the amount of gene flow ( Nm ) at several levels. Gene flow among British populations was extremely high ( Nm = 36.51), one of the highest estimates reported for plants. Among eight European populations gene flow was lower (but still considered high) at Nm = 2.47. Trans-Atlantic gene flow was low ( Nm = 0.0926).
F -statistics were used to assess population heterogeneity and substructuring. The data indicate that, compared with other species, there is very little genetic differentiation among British populations of bracken. Indeed, it appears that the whole island is behaving as a single randommating population. This result is consistent with high levels of gene flow. Only one population (on the Isle of Arran) showed statistically significant genetic substructuring. Habitat heterogeneity on the island and age structure are hypothesized as possible causes of this result.
The data reported here support previous studies demonstrating that bracken is genetically polymorphic and is an outcrossing species.     

Evaluation of physiological traits for improving drought tolerance in faba bean (<Emphasis Type="Italic">Vicia faba</Emphasis> L.)

Among grain legumes, faba bean is becoming increasingly popular in European agriculture due to recent economic and environmental interests. Faba bean can be a highly productive crop, but it is sensitive to drought stress and yields can vary considerably from season to season. Understanding the physiological basis of drought tolerance would indicate traits that can be used as indirect selection criteria for the development of cultivars adapted to drought conditions. To assess genotypic variation in physiological traits associated with drought tolerance in faba bean and to determine relationships among these attributes, two pot experiments were established in a growth chamber using genetic materials that had previously been screened for drought response in the field. Nine inbred lines of diverse genetic backgrounds were tested under adequate water supply and limited water conditions. The genotypes showed substantial variation in shoot dry matter, water use, stomatal conductance, leaf temperature, transpiration efficiency, carbon isotope discrimination (Δ¹³C), relative water content (RWC) and osmotic potential, determined at pre-flowering vegetative stage. Moisture deficits decreased water usage and consequently shoot dry matter production. RWC, osmotic potential, stomatal conductance and Δ¹³C were lower, whereas leaf temperature and transpiration efficiency were higher in stressed plants, probably due to restricted transpirational cooling induced by stomatal closure. Furthermore, differences in stomatal conductance, leaf temperature, Δ¹³C and transpiration efficiency characterized genotypes that were physiologically more adapted to water deficit conditions. Correlation analysis also showed relatively strong relationships among these variables under well watered conditions. The drought tolerant genotypes, ILB-938/2 and Melodie showed lower stomatal conductance associated with warmer leaves, whereas higher stomatal conductance and cooler leaves were observed in sensitive lines (332/2/91/015/1 and Aurora/1). The lower value of Δ¹³C coupled with higher transpiration efficiency in ILB-938/2, relative to sensitive lines (Aurora/1 and Condor/3), is indeed a desirable characteristic for water-limited environments. Finally, the results showed that stomatal conductance, leaf temperature and Δ¹³C are promising physiological indicators for drought tolerance in faba bean. These variables could be measured in pot-grown plants at adequate water supply and may serve as indirect selection criteria to pre-screen genotypes.     

The effects of nitrogen stress on the stable carbon isotope composition, productivity and water use efficiency of white spruce (Picea glauca (Moench) Voss) seedlings

A, assimilation rate
a, fractionation against ¹³C for CO₂ diffusion through air
b, net fractionation against ¹³C during CO₂ fixation
C_a, ambient CO₂ concentration
C_c, CO₂ concentration at the chloroplast
C_i, intercellular CO₂ concentration
D, vapour pressure deficit
E_n, needle transpiration rate
E_p, whole plant water use
g_w, leaf internal transfer conductance to CO₂
g_s, stomatal conductance to water vapour
L, projected leaf area
NUE, nitrogen use efficiency
PEP, phosphoenolpyruvate
Rubisco, ribulose-1,5-biphosphate carboxylase
TDR, time domain reflectometry
WUE, water use efficiency
Δ, carbon isotope discrimination
δ¹³C, carbon isotope abundance parameter
δ¹³C_a, carbon isotopic composition of atmospheric CO₂
θ, volumetric soil water content

The effect of nitrogen stress on needle δ¹³C, water-use efficiency (WUE) and biomass production in irrigated and dry land white spruce (Picea glauca (Moench) Voss) seedlings was investigated. Sixteen hundred seedlings, representing 10 controlled crosses, were planted in the field in individual buried sand-filled cylinders. Two nitrogen treatments were imposed, nitrogen stressed and fertilized. The ranking of δ¹³C of the crosses was maintained across all combinations of water and nitrogen treatments and there was not a significant genetic versus environmental interaction. The positive relationships between needle δ¹³C, WUE and dry matter production demonstrate that it should be possible to use δ¹³C as a surrogate for WUE, and to select for increased WUE without compromising yield, even in nitrogen deficient environments. Nitrogen stressed seedlings had the lowest needle δ¹³C in both irrigated and dry land conditions. There was a positive correlation between needle nitrogen content and δ¹³C that was likely associated with increased photosynthetic capacity. There was some indication that decreased nitrogen supply led to increased stomatal conductance and hence lower WUE. There was a negative correlation between intrinsic water use efficiency and photosynthetic nitrogen use efficiency (NUE). This suggests that white spruce seedlings have the ability to maximize NUE when water becomes limited. There was significant genetic variation in NUE that was maintained across treatments. Our results suggest that in white spruce, there is no detectable effect of anaplerotic carbon fixation and that it is more appropriate to use a value of 29‰ (‘Rubisco only’) for the net discrimination against ¹³C during CO₂ fixation. This leads to excellent correspondence between values of C_i/C_a derived from gas exchange measurements or from δ¹³C.     

Crop growth,water-use efficiency and carbon isotope discrimination in groundnut (Arachis hypogaea L.) genotypes under end-of season drought conditions*

Ten groundnut genotypes were grown under adequately irrigated conditions or subjected to drought during the pod filling phase (83–113 days after sowing) in a medium deep Alfisol at the ICRISAT Centre during the 1986–1987 post-rainy season. Crop growth was measured in both treatments, but transpiration (7) and water-use efficiency (W) were quantified only in the drought treatment. Leaf samples from both treatments were assayed for discrimination against ¹³CO₂ fixed in leaves (Δ) to examine the relationships between Δ, crop growth, and W under field conditions. The shoot dry matter accumulated during the period of drought (Y) ranged from 72–150 g m^-2 and was closely related to transpiration. This indicates scope for selection of traits and practices to increase T. Water-use efficiencies ranged from 1.38–2.50 g kg^-1 and were inversely related to Δ in eight out of the 10 genotypes. For the other two genotypes, there was evidence that T was underestimated by field measurements. Water-use efficiency and transpiration were not correlated suggesting that these two traits might be combined through breeding. Variation between genotypes was greatest for the partitioning of total dry matter to pods (73%), followed by water-use efficiency (31%) and transpiration (29%). Crop growth rates were negatively related to Δ under irrigated conditions but not under drought.     

Photosynthesis on individual leaves of sugar beet (Beta vulgaris) during the ontogeny at variable water regimes

It is well known that the extent of yield reduction depends not only on the severity of water stress but also on the stage of plant development. Assessing photosynthetic response of individual leaves to water deficit during the ontogeny may, therefore, offer a clue to better understand the whole plant behaviour. This research aimed at investigating the influence of early and late water stress on net photosynthesis (P_n), carbon‐isotope discrimination and other related traits on individual leaves during ontogeny. Sugar beet plants were grown in rain‐sheltered soil columns of relevant volume (300 L), subdivided into well‐watered (WW); early (S1) and late (S2) stress. In general, water stress significantly reduced leaf lifespan and P_n. Relieving the stress at about one‐third and two‐thirds of potential leaf life substantially restored P_n at the levels of WW. Stressing a previously WW leaf brought about a comparatively heavier loss than stressing a leaf since the beginning. As for leaves at different phenological times, the early leaves had higher initial photosynthetic peaks but steeper falls during their lives. An insight into the relationships between P_n and substomatal CO₂ concentration (C_i) shows that in mature leaves the photosynthetic restoration following stress relief did not entail a full recovery of the electron transport rate, the parameter most severely affected by the stress. The partial reversibility of the effects of water deficiency, associated to the anticipated leaf senescence and to the natural slow‐down of net assimilation during leaf life, may be seen as a key factor in predicting to what extent the plant can tolerate drought and the damages caused by water stress.     

Water resource partitioning,stem xylem hydraulic properties,and plant water use strategies in a seasonally dry riparian tropical rainforest

This study investigated seasonal variation in the origin of water used by plants in a riparian tropical rainforest community and explored linkages between plant water source, plant xylem hydraulic conductivity and response to the onset of dry conditions. The study focused on five co-dominant canopy species, comprising three tree species (Doryphora aromatica, Argyrodendron trifoliolatum, Castanospora alphandii) and two climbing palms (Calamus australis and Calamus caryotoides). Stable isotope ratios of oxygen in water (¹⁸O) from soil, groundwater, stream water and plant xylem measured in the wet season and the subsequent dry season revealed water resource partitioning between species in the dry season. Measurement of stem-area-specific hydraulic conductivity (K_S) in the wet season and subsequent dry season showed a significant dry-season loss of K_S in three of the five species (Castanospora alphandii, Calamus australis and C. caryotoides) and a decrease in mean K_S for all species. This loss of hydraulic conductivity was positively correlated with the difference between wet-season and dry-season midday leaf water potentials and with leaf carbon isotope discrimination, indicating that plants that were less susceptible to loss of conductivity had greater control over transpiration rate and were more water-use efficient.     

Effects of enhanced UVB on populations of the phloem feeding insect Strophingia ericae (Homoptera: Psylloidea) on heather (Calluna vulgaris)

Heather plants (Calluna vulgaris (L.) Hull) were grown for two years (April 1994–October 1996) under ambient or enhanced ultraviolet-B radiation (UVB: 280–315 nm), provided as a modulated treatment simulating a 15% ozone depletion (seasonally adjusted). Populations of the psyllid (Strophingia ericae (Curtis)) were measured before treatment and at yearly intervals thereafter. Before treatment there was no significant difference in the psyllid populations between treatments, or between the experimental and source populations. Enhanced UVB progressively produced a reduction in S. ericae populations compared with controls over 27 months. Analyses of C, N, total water soluble phenolics, total free amino acids and measurements of leaf angles and distances between leaflets demonstrated no effects of UVB treatment. However, concentrations of the amino acid isoleucine were lower (28%) in C. vulgaris exposed to the enhanced UVB treatment. Over the duration of the experiment the psyllid population structure at Lancaster changed from that typical of the upland site of origin (two-year cycle with overlapping cohorts) to a one-year life cycle typical of lowland sites, but this was independent of UVB treatments. Reduced isoleucine might explain the negative effects of elevated UVB on psyllid population numbers, but the precise effects of UVB on host chemistry and morphology are unknown. The problem of interpreting herbivore responses to enhanced UVB treatments in the field is discussed.     

Genotypic variations in carbon isotope discrimination, transpiration efficiency, and biomass production in rice as affected by soil water conditions and N

Genotypic and environmental (soil water regime and N level) variation in carbon isotope discrimination (CID) in relation to the gas exchange, transpiration efficiency (A/T), and biomass production were investigated in field experiments using eleven rice (Oryza sativa L.) genotypes. The results showed that genotype was more dominant for variation in CID than in total biomass. Genotypic ranking in CID was consistent across environments because of small genotype × environment interactions. Japonica genotypes tended to have lower CID than indica genotypes. Higher soil water and lower N rate significantly increased CID. Variation in CID was slightly smaller for water regime than for genotype. There was a negative correlation between CID andA/T among genotypes within water regimes. Genotypic variation in CID was associated mainly with variation in stomatal conductance under all soil water regimes and with photosynthetic capacity in late growth stages under aerobic soil conditions. The decrease in CID at higher N was probably due to lower stomatal conductance under aerobic soil conditions and to higher photosynthetic rates under submerged soil conditions. The correlation between biomass and CID was not clear in aerobic soil, whereas it was positive in submerged soil, which indicated that the significance of lower or higher CID for improving biomass productivity may differ under different soil water regimes. Overall, the results implied a possible use of CID as a selection criterion for genotypic improvement inA/T and productivity in rice.     

土壤生物多样性对植物利用营养物质和水分的影响

土壤生物多样性影响土壤营养物质和水等自然资源的有效利用是农业生态系统的重要功能.有证据表明土壤微生物多样性可提高土壤营养物质和水的利用率.土壤动物对土壤营养物质和水的有效利用也产生了明显的影响,它主要是通过促进营养物质的转化和改善土壤水渗透率而间接实现的.从已有文献的报道来看,单从某一个方面来研究土壤生物多样性对土壤营养物质和水的有效利用的影响并不能全面地反应它们之间相互作用的规律.因此未来的研究应当把土壤生物多样性、土壤营养物质和水的供给与植物生物多样性、植物营养物质和水的利用率结合起来,这样才有可能从理论上对土壤生物多样性与营养物质和水的利用之间的关系有更加深刻的认识并运用到实践中.