Drought survival in Dactylis glomerata and Festuca arundinacea under similar rooting conditions in tubes

Drought survival in perennial forage plants involves different adaptative responses such as delay of dehydration through water uptake, limitation of water loss and tolerance of tissues to dessication. To compare the importance of these responses in contrasting cultivars of forage grasses at the whole plant level, we carried out two experiments under glasshouse conditions. Plants of cocksfoot (Dactylis glomerata L.) cultivars, cvs. Currie, Medly (both of Mediterranean origin) and Lutetia (of continental origin), and of tall fescue (Festuca arundinacea L.) cv. Centurion (Mediterranean) were grown in 60 cm-deep cylinders to eliminate the effect of differences of root depth on water availability whilst allowing severe drought to be imposed at a realistic rate. In both experiments, the cvs. were ranked similarly for plant survival, with high mortality for Centurion, low for the Mediterranean cocksfoots Currie and Medly, and intermediate for Lutetia. These differences could not be ascribed to water use during most of the drought period since water uptake and decrease in leaf extension were not significantly different between species and cultivars. However, resistant cvs. of cocksfoot were able to extract water for a longer period and at a lower soil water potential (s) than other cvs. The critical s at plant death was –3.8 and –3.6 MPa for Medly and Currie and –3.0-,–2.6 MPa for Lutetia and Centurion. Moreover, at a low soil water reserve (15–2%), membrane stability and water content were maintained for longer in enclosed immature leaf bases of cocksfoots cultivars, whereas the fescue Centurion exhibited accelerated lamina senescence and steady increase of membrane damage in surviving tissues. Therefore, it is proposed that the drought resistance of tall fescue in the field can mainly be ascribed to its ability to develop a deep root system. In cocksfoot, dehydration tolerance in surviving tissues and the ability of roots to extract water at low soil water potentials may, in addition to root depth, contribute significantly to plant survival under severe drought.     

Nitrate uptake ability by maize roots during and after drought stress

The effects of different intensities and durations of soil drought and re-watering on the nitrate uptake ability of maize roots were studied. Plants were grown in split-root containers with one part of the root system subjected to different intensities and durations of soil drought and re-watering while the other part of the root system was continuously watered to 23% (w/w) soil water content (70% water capacity). Experiments were performed in split-root containers to maintain a high growth rate, thus ensuring high nutrient demand of the shoot irrespective of the soil water regime. To avoid limitation of nitrate uptake by transport processes in the dry soil, and to ensure a uniform ¹⁴N/¹⁵N ratio at the root surface, ¹⁵N was applied to the roots by placing them into an aerated nutrient solution with 0.5 mM Ca(¹⁵NO₃)₂. Shoot elongation and biomass were only slightly affected by drought in one root compartment when the soil in the other root compartment was kept wet. Therefore, the growth-related nutrient demand of the shoot remained at a high level. At moderate levels of soil drought (10% w/w water content) the ability of the roots for N-uptake was not affected even after 10 d of drought. N-uptake ability was reduced to about 20% of the well-watered control only when the soil water content was decreased to 5%. Total soluble sugar content of the roots increased with increasing soil drought, indicating that low N-uptake ability of roots subjected to severe soil drought was not caused by low assimilate supply from the shoot. Nitrate uptake ability of roots maintained in very dry soil (5% soil water content w/w) even for a prolonged period of 8 d, recovered within 3 d following re-watering. Root growth increased one day after re-watering. A short-term experiment with excised roots formerly subjected to severe soil drought showed that nitrate uptake ability recovered in old and young root segments after 2 d of re-watering. Obviously, the increase in N-uptake ability after re-watering was caused not only by new root growth but also by recovery of the uptake ability of formerly stressed roots.     

Water Use Characteristics of Dactylis glomerata L., Lolium perenne L. and L. multiflorum Lam. Plants

Cocksfoot (Dactylis glomerata L.). perennial ryegrass (Loliumperenne L.) and Italian ryegrass (L. multiflorum Lam.) plantswere grown on deep (75–95 cm) columns of soil in glasshousesand growth rooms with and without irrigation. The species inwhich growth declined least rapidly after water had been withheldwere those which transpired most slowly. During early establishmentin the glasshouse cocksfoot transpired least because of slowroot growth. In the growth room, when root systems were deeperand denser, perennial ryegrass transpired least because of lowleaf water conductance. Results are discussed in relation to(a) drought resistance in the three species; (b) breeding forincreased drought resistance through modifying root distributionand leaf water conductance; and (c) the use of isolated soilcolumns in water relations studies. Dactylis glomerata L., Lolium perenne L., Lolium multiflorum Lam., cocksfoot, perennial ryegrass, Italian ryegrass, transpiration, roots, leaf water conductance     

Effectiveness of phytoremediation as a secondary treatment for polycyclic aromatic hydrocarbons (PAHs) in composted soil

A greenhouse study was conducted over a 12-month period to investigate the fate of polycyclic aromatic hydrocarbons (PAHs) in soil using phytoremediation as a secondary treatment. The soil was pretreated by composting for 12 weeks, then planted with tall fescue (Festuca arundinacea), annual ryegrass (Lolium multiflorum), and yellow sweet clover (Melilotus officinalis). Two sets of unvegetated controls also were evaluated, one fertilized and one unfertilized. Total PAH concentrations decreased in the tall fescue, annual ryegrass, and yellow sweet clover treatments by 23.9%, 15.3%, and 9.1%, respectively, whereas the control was reduced by less than 5%. The smaller two- and most of the three-ringed compounds--naphthalene, acenaphthylene, acenaphthene, fluorene, and anthracene--were not found in detectable concentrations in any of the treatments. The most probable number analysis for microbial PAH degraders did not show any statistically significant differences among treatments. There were significant differences among treatments (p < 0.05) for the residual concentrations of five of the target PAHs. Root surface area measurements indicated that tall fescue and annual ryegrass both had significantly higher root surface area than yellow sweet clover, although the two species were not significantly different from each other. The tall fescue treatment resulted in the highest root and shoot biomass, followed by annual ryegrass and yellow sweet clover, and also had the highest percent of contaminant removal after 12 months. These results imply a positive relationship between plant biomass development and PAH biodegradation.     

Effects of tall fescue turf on growth and nitrogen fixation potential of the woody legume Lupinus albifrons

The effect of tall fescue turf on growth, flowering, nodulation, and nitrogen fixing potential of Lupinus albifrons Benth. was examined for greenhouse and field grown plants. No allelopathic effect was observed for lupine plants treated with tall fescue leachates. The nitrogen-fixing potential measured by nodule dry weight and acetylene reduction rates was not significantly affected by tall fescue turf.Both the greenhouse and field studies showed that the growth, sexual reproductive allocation and number of inflorescences were significantly reduced when lupine plants were grown with tall fescue. The root-length densities of tall fescue turf and lupine monoculture were measured. The tall fescue turf had 20 times higher root-length density (20 cm cm^-3 soil) than the lupine plant monoculture. This suggests that intense competition at the root zone may be a dominant factor which limits the growth of the lupine plants.The flowering characters of the lupine plants were improved by phosphorus fertilization. Transplanting of older lupine plants into the turf substantially alleviated the tall fescue turf competitive effect.     

Effect of plant species on the larvae of gastrointestinal nematodes which parasitise sheep

Faeces containing Trichostrongylus colubriformis and/or Ostertagia circumcincta eggs were used to provide four contaminations in each of 2 years on plots of browntop, Yorkshire fog, ryegrass, tall fescue, lucerne, chicory, cocksfoot, white clover, and prairie grass and in the second year a mixed sward of ryegrass/white clover. Third stage larvae were recovered from faeces and from four strata of herbage, 0–2.5, 2.5–5, 5–7.5 and >7.5 cm above the soil surface at 2, 4, 6, 8, 11, and 14 weeks after faeces were deposited on the swards. Herbage species had a significant (P &lt; 0.0001) effect on the number of larvae recovered. Greatest numbers of larvae, as indicated by ranking analysis, were recovered from Yorkshire fog, ryegrass, and cocksfoot and lowest numbers from white clover and lucerne. The difference between herbages in numbers of larvae recovered was due to the ‘‘development success’’, the ability of larvae to develop to the infective stage and migrate on to herbage, rather than ‘‘survival’’, the rate of population decline once on the herbage. Faecal degradation was most rapid from white clover and browntop, intermediate from tall fescue, lucerne, prairie grass, cocksfoot, and ryegrass, and slowest from Yorkshire fog swards. The numbers of larvae recovered from herbages were related (r² = 0.59, P &lt; 0.05) with the faecal mass remaining. A greater proportion of the total larvae recovered from the herbage was recovered from the bottom stratum of Yorkshire fog and prairie grass than from white clover, with the other herbages intermediate, indicating that larvae had greater difficulty migrating up Yorkshire fog and prairie grass than the other herbage species. In most herbage species, despite more larvae being recovered from the lowest stratum, larval density (L₃/kg herbage DM) was highest in the top stratum. This study has demonstrated that herbage species can have a significant impact on the population dynamics and vertical migration of T. colubriformis and O. circumcincta larvae.     

Photosynthesis,respiration, and carbon allocation of two cool-season perennial grasses in response to surface soil drying

The study was conducted to investigate carbon metabolic responses to surface soil drying for cool-season grasses. Kentucky bluegrass (Poa pratensis L.) and tall fescue (Festuca arundinaceae Schreb.) were grown in a greenhouse in split tubes consisting of two sections. Plants were subjected to three soil moisture regimes: (1) well-watered control; (2) drying of upper 20-cm soil (upper drying); and (3) drying of whole 40-cm soil profile (full drying). Upper drying for 30 d had no dramatic effects on leaf water potential (Ψ_leaf) and canopy photosynthetic rate (P_n) in either grass species compared to the well-watered control, but it reduced canopy respiration rate (R_canopy) and root respiration rate in the top 20 cm of soil (R_top). For both species in the lower 20 cm of wet soil, root respiration rates (R_bottom) were similar to the control levels, and carbon allocation to roots increased with the upper soil drying, particularly for tall fescue. The proportion of roots decreased in the 0-20 cm drying soil, but increased in the lower 20 cm wet soil for both grass species; the increase was greater for tall fescue. The Ψ_leaf, P_n, R_canopy, R_top, R_bottom, and carbon allocation to roots in both soil layers were all significantly higher for upper dried plants than for fully dried plants of both grass species. The reductions in R_canopy and R_top in surface drying soil and increases in root respiration and carbon allocation to roots in lower wet soil could help these grasses cope with surface-soil drought stress. This revised version was published online in June 2006 with corrections to the Cover Date.     

Common and newly-recorded forage crop diseases in Ireland

Surveys of forage crop diseases in the Republic of Ireland have been conducted annually since 1965. Over fifty fungal diseases have been noted, about half of which are of likely economic importance. Disease control investigations were carried out on the more prevalent diseases, such as crown rust and Ophiobolus patch of ryegrass, leaf fleck of cocksfoot and powdery mildew of tall fescue. Some pathogens previously unrecorded in the British Isles were found. These included Spermospora lolii and Lolium mottle virus on ryegrass, and Mastigosporium kitzebergense on timothy.     

The nitrogen handling characteristics of white clover (Trifolium repens L.) cultivars and a perennial ryegrass (Lolium perenne L.) cultivar

Ryegrass (Lolium perenne L.) and white clover (Trifolium repens L.) have contrasting responses to soil mineral N availability and clover has the ability to fix atmospheric N(2) symbiotically. It has been hypothesized that these differences are the key to understanding grass-clover coexistence and vegetative dynamics in pastures. However, the whole plant response of clover and ryegrass to mineral N availability has not been fully characterized and inter-cultivar variability in the N-handling dynamics of clover has not been assessed. A detailed experimental study to address these issues was undertaken. For all clover cultivars and ryegrass, mass specific mineral N uptake rates (of whole plants) were similar saturating functions of mineral N availability. For all clover cultivars total N assimilation rates, whole plant C : N ratios and root : shoot ratios were independent of mineral N availability. Clover growth rates were also independent of mineral N availability except for a slight (<10%) reduction at very low N availability levels. Specific N(2) fixation rate (whole plant) was precisely controlled to ensure fixation balanced the deficit between mineral N uptake and the total N assimilation required to maintain constant whole plant C : N ratio. There was always a deficit between N uptake and the total N assimilation required to maintain C : N ratio. Consequently, some N(2) fixation remained engaged even at high mineral N availability levels. All inter-cultivar variation in N(2) fixation dynamics could be attributed to variations in growth rate. Clover mass specific growth rate declined as plant size increased. Ryegrass specific growth rate, whole plant C : N ratio and root : shoot ratio were dependent on N availability. Increased N availability led to increased growth rate and decreased C : N and root : shoot ratios. Specific growth rate was also dependent on plant size, growth rate declining as plant size increased. It is concluded that clover inter-cultivar variation in field performance is unlikely to be a consequence of variation in N-handling characteristics. Inter-cultivar differences in growth rate are likely to be a much more important source of variation.     

FIELD EXPERIMENTS ON CEREAL ROOT EELWORM

In 1955-59 experiments were carried out in south-west England on field sites heavily infested by the cereal root eelworm, to determine the behaviour of the soil population under various crops.
Under three common types of ley, the population fell to less than 5% of its original level in 3 years.
Of six pure grass stands tested, the population fell most sharply under cocksfoot and tall fescue.
Observation plots under the four cereals illustrate their different efficiencies as hosts.     

Comparison of tall fescue (Cyperales: Gramineae) to other cool-season turfgrasses for tolerance to European chafer (Coleoptera: Scarabaeidae)

Three cultivars of tall fescue, Festuca arundinacea Schreb., were compared with three cultivars each of fine fescue (Festuca spp.), Kentucky bluegrass (Poa pratensis L.), and perennial ryegrass (Lolium perenne L.) to evaluate tolerance to root-feeding by European chafer grubs, Rhizotrogus majalis (Razoumowsky). Potted turfgrasses were infested with initial densities equivalent to 33 or 66 grubs per 0.1 m2 on 19 August 2000. More grubs were added in late September and October, bringing the total to 66 or 143 grubs per 0.1 m2. Plant growth, root loss, weight gain, and survival of grubs were measured. The experiment was repeated in fall of 2001 with an initial density of 66 grubs per 0.1 m2. The proportion of root mass lost as a result of grub feeding was a function of turf species, root growth, grub survival, and grub growth during the test. Grubs gained the most weight and consumed the most roots when feeding on fine fescue. Fine fescue suffered the greatest percentage of root loss in 2000, despite having the most rapid root growth and largest mass in control pots. Cultivars of tall fescue appeared to be the most tolerant of grub feeding, having the smallest reduction in root mass in both years. Data from fine fescue, Kentucky bluegrass, and perennial ryegrass cultivars were not as consistent as tall fescue, because for some cultivars root growth and grub survival were different between years. We also found that grubs increased in mass by 20% when the mass of available roots was doubled.     

Endophytic fungus improves growth and metal uptake of Lolium arundinaceum Darbyshire ex. Schreb

The effect of endophyte infection on plant growth, cadmium (Cd) uptake, and Cd translocation was investigated using tall fescue (Lolium arundinaceum) grown in greenhouses in contaminated solution. Endophyte infection significantly increased tiller number and biomass of the host grass under both control and Cd-stress conditions. Endophyte infection not only enhanced Cd accumulation in tall fescue, but also improved Cd transport from the root to the shoot. Under 20 mg L(-1) Cd stress, the phytoextraction efficiency of endophyte-infected (EI) tall fescue was 2.41-fold higher than endophyte-free plants. Although the total Cd accumulation in EI tall fescue was insufficient for practical phytoextraction applications, the observed high biomass production and tolerance of stress from abiotic factors including heavy metals, gives endophyte/plant associations the potential to be a model for endophyte-assisted phytoremediation of metal-polluted soils.     

Effect of different soil and nitrogen fertilizer conditions on 32P-labelled phosphate uptake by two grass species

In two years of trials, roots of ryegrass took up more ³²P-labelled phosphate than roots of fescue. Application of 672 kg N ha^-1 increased phosphate absorption compared with application of 112 kg N ha^-1. Roots in mineral soil absorbed more phosphate than those in peat soil. In both soils uptake decreased as depth of phosphate injection increased from 5 to 30 cm. An interaction occurred whereby roots in the intermediate depth (10–22-5 cm) in peat absorbed less phosphate than in mineral soil and this was apparently unrelated to the exchange or sorption properties of the soil.     

Inter-clonal difference in sodium uptake within the speciesphleum pratense andfestuca pratensis

Summary A study was made of intra-species differences in sodium uptake by timothy (Phleum pratense) and meadow fescue (Festuca pratensis), two pasture grasses that are generally known as being low in Na-content compared with perennical ryegrass (Lolium perenne) and cocksfoot (Dactylis glomerata). Na²² was used with NaNO₃ as a tracer in a fertilizer experiment on seedlings grown on trays placed on benches in the greenhouse. With timothy significant clonal differences in Na-uptake varied from one half to the double, but the Na-contents were still appreciably lower than in ryegrass or cocksfoot.     

Involvement of antioxidants and lipid peroxidation in the adaptation of two cool-season grasses to localized drought stress

In natural environments, drought often occurs in surface soil while water is available for plant uptake deeper in the soil profile. The objective of the study was to examine the involvement of antioxidant metabolism and lipid peroxidation in the responses of two cool-season grasses to surface soil drying. Kentucky bluegrass (Poa pratensis L) and tall fescue (Festuca arundinacea Schreb.) were grown in split tubes, consisting of two sections (each 10 cm in diameter and 20 cm long). Grasses were subjected to three soil moisture regimes: (a) well-watered control: whole soil profile was watered; (b) surface drying: surface 20 cm of soil was dried by withholding irrigation and the lower 20 cm of soil was watered; (c) full drying: whole soil profile was dried. Surface drying had no effects on relative water content (RWC) and chlorophyll content (Chl) for both grasses and only slightly reduced shoot growth for tall fescue. Superoxide dismutase (SOD) activity increased, while catalase (CAT) and peroxidase (POD) activities remained unchanged during most periods of surface drying. Malondialdehyde (MDA) content was unaffected by surface drying for tall fescue, but increased initially and then decreased to the control level for Kentucky bluegrass. Under full drying, RWC, Chl content, and shoot dry weight decreased, but MDA content increased in both grasses; SOD and POD activities initially increased transiently and then decreased; CAT remained unchanged for 25 days and then decreased. These results suggested that both Kentucky bluegrass and tall fescue were capable of surviving surface soil drying. This capability could be related to increases in antioxidant activities, particularly SOD and CAT. However, full drying suppressed antioxidant activities and induced lipid peroxidation.     

Nutritive value of forage biomass from sainfoin mixtures

Forage quality characteristics of field-grown mixtures of sainfoin with cocksfoot (50:50%), sainfoin with tall fescue (50:50%), and the same with the addition of subterranean clover in their composition (33:33:33%) were measured. Forage biomass from the mixtures of sainfoin with cocksfoot had generally higher forage quality than mixtures with tall fescue. It had higher crude protein content (11.52% of dry matter (with 1.07% units), significantly higher digestibility (61.74%) (with 6.51% units), higher neutral detergent fiber content (53.42%) (with 3.22% units), higher nutritive value (Unité Fourragère Viande – Unité Fourragère Lait, 0.690–0.583) and higher protein feeding value (Total Digestible Protein – Protein digestible dans l’intestine in dependence of nitrogen – Protein digestible dans l’intestine in dependence of energy), 72–70–79 g/kg of dry matter. Forage biomass showed more balanced basic chemical composition after the addition of subterranean clover, i.e.: higher crude protein content (with 0.30% units) and lower crude fiber content (with 0.14% units) for mixtures with cocksfoot; higher digestibility (with 0.29% units) for mixtures with cocksfoot; lower neutral detergent fiber content (with 0.45% units) for mixtures with cocksfoot and with 3.15% units for mixtures with tall fescue, higher energy feeding value (Unité Fourragère Viande – Unité Fourragère Lait) (with 0.007–0.012 for mixtures with cocksfoot and with 0.009–0.014 for mixtures with tall fescue), higher protein feeding value for both mixtures with cocksfoot and tall fescue. Forage biomass from mixtures of sainfoin with cocksfoot and Trifolium subterraneum ssp. brachycalicinum had the highest crude protein (11.89% of dry matter), the lowest crude fiber content (27.07% of dry matter) and the highest digestibility (62.81% of dry matter).     

Survival and recovery of perennial forage grasses under prolonged Mediterranean drought: I. Growth, death, water relations and solute content in herbage and stubble

Swards of Dactylis glomerata cultivars (cvs) KM2 and Lutetia and of Lolium perenne cvs Aurora and Vigor were grown under full irrigation or prolonged summer drought (80 d) in a field experiment in the South of France.
After irrigation was withheld, leaf extension rates of all cvs fell by 90% within 9–12 d, and rapid scorching of laminae followed. Tiller mortality at the end of the drought was very different in the cocksfoot cvs (4% for KM2 and 76% for Lutetia) and intermediate (41%) for both ryegrass cvs. Following re-watering, rates of herbage regrowth were closely correlated with tiller survival. Measured minerals contributed c . 0·52 MPa to osmotic potential in all treatments, whereas water-soluble carbohydrates (WSC) contributed 0·25 MPa under irrigation and 0·46 MPa during drought.
There was no systematic difference between the two species for summer survival under severe drought, but large differences between the cocksfoot cvs. The traits most strongly associated with superior survival were: (a) a deep root system and greater water uptake at depth; (b) low water and osmotic potentials in surviving laminae, i.e. better tolerance to dehydration; (c) large pool-size of WSC reserves (fructans having degree of polymerization >4) in entire tiller bases (stubble); (d) low accumulation of proline in stubble; (e) rapid nitrogen uptake after rewatering.     

Grass species and their fungal symbionts affect subsequent forage growth

Persistence of forage grasses is enhanced through the deliberate and selective use of symbiotic fungal endophytes that confer benefits, particularly pest resistance. However, they have also been implicated in reduced plant community diversity as a result of directly or indirectly enhancing competitive ability. A relatively underexplored mechanism by which endophytes might influence pasture plant composition is by altering the biotic or abiotic soil conditions. To examine the soil conditioning effects of forage grass species and their fungal symbionts we tested the responses of three pasture plants, perennial ryegrass, prairie grass, and white clover in nine different soils that had been conditioned by monocultures of endophyte-containing (E+), or endophyte-free (E?), perennial ryegrass, tall fescue, or meadow fescue. Conditioning grass species had little effect on the responses of perennial ryegrass and prairie grass regardless of E+ or E? treatments. In contrast, conditioning species had a strong effect on the response of white clover, resulting in reduced biomass when grown in perennial ryegrass conditioned soils. The presence of endophyte also had significant growth consequences for white clover, but was either positive or negative depending on the conditioning grass species. In comparison to their respective E? treatments, E+ tall and meadow fescue conditioned soils resulted in reduced biomass of white clover, whereas E+ perennial ryegrass conditioned soils resulted in increased biomass of white clover. Among the conditioning strains (AR1, AR37, NEA2, WE) of E+ perennial ryegrass, white clover showed significantly different responses, but all responses were positive in comparison to the E? treatment. By examining the effects of several grass species and endophyte strains, we were able to determine the relative importance of grass species vs. fungal symbiont on soil conditioning. Overall, the conditioning effect of grass species was stronger than the effects associated with endophyte, particularly with regard to the response of white clover. We conclude that both grass species and their fungal endophytes can influence pasture plant community composition through plant–soil feedback.     

超声波对3种老化牧草种子萌发和幼苗生长的影响

利用四因素(超声时间、超声温度、输出功率和浸种时间)三水平[L₉(3⁴)]正交试验,对自然老化的高羊茅(在室温条件下储藏1年和5年)、黑麦草(储藏5年)和新麦草(储藏6年)种子进行超声波处理,研究超声波对老化种子萌发、幼苗生长的作用及其生理机制.结果表明: 超声温度是影响老化种子萌发的主要因素.适宜的超声波处理可以提高老化的高羊茅(储藏5年)和新麦草种子发芽率,显著促进胚根和胚芽伸长,而对老化的高羊茅(储藏1年)和黑麦草种子萌发无明显改善,但可促进其胚根伸长.超声波可通过提高超氧化物歧化酶、过氧化物酶活性及显著降低丙二醛含量来恢复种子活力.超声波处理效果存在参数阈值效应,通过回归模型分析得出其处理老化草种的最优参数为:超声时间22 min,超声温度26 ℃,超声输出功率254.29 W,浸种时间2.89 h.     

Long-term drought stress alters nitrogenase activity and carbon translocation in split-root cultured Alnus incana

Split-root cultured grey alder, Alnus incana (L.) Moench., was grown in sand in cuvettes with a continuous supply of nutrient solution. During the drought treatment for up to 9 days the supply of solution was withheld from one of the split-root halves. After 2–3 days of treatment, soil water became depleted and the unwatered root halves were at a constant drought stress, water potential (Ψ_nodules) = -1.1 to -1.6 MPa. Nitrogenase activity in the dry half decreased to about 70% of the initial value during the first 2–3 days and then stayed at this level. The water supply to the shoot from the wet root half was high and only a temporary slight decrease in photosynthesis and stomatal conductance was found in drought-stressed split-root plants. Labelling studies showed a reduced translocation of photoassimilates to the dry nodules. The fixation of CO₂ in the nodules seemed to be more tolerant to drought than nitrogenase activity. During the drought treatment there was an osmotic adjustment from -0.9 to -1.7 MPa, but no change in the storage of starch in the nodules. In alders where parts of the root system is kept dry these roots acclimate and continue a persistent nitrogenase activity.