Hydraulic conductance and soil water potential at the soil-root interface of Pinus pinaster seedlings inoculated with different dikaryons of Pisolithus sp

Seedlings of maritime pine (Pinus pinaster Ait.) were inoculated with different dikaryons of Pisolithus sp. from South Africa to determine the influence of extension of the extramatrical phase and diameter of the mycelial strands on water relations parameters including xylem water potential (Psi(x)), soil water potential at the soil-root interface (Psi(s)) and hydraulic conductance (L(p)) during and after a period of water stress. Seedlings inoculated with dikaryons having an extensive extramatrical phase and large diameter mycelial strands showed higher Psi(s) (-2 MPa) during severe water stress than seedlings inoculated with dikaryons producing fine hyphae and sparse extramatrical phases (-3.8 MPa). Seedlings inoculated with strand-forming dikaryons recovered faster from water stress than did non-inoculated seedlings or seedlings inoculated with non-strand-forming dikaryons. Architectural aspects of the extramatrical phase, including the presence of large diameter mycelial strands or fine hyphae, influenced the soil-root contact and the water relations of an inoculated host plant. When water stress was not limiting, the architecture of the extramatrical phase did not have a large effect on Psi(s). It is suggested that the architecture of the extramatrical phase influences the resistance to water flow through the soil-root interface and that large mycelial strands increase the water flow by bridging the gap between the soil and the root. These changes in physiology indicate that dikaryons can improve the survival of Pinus pinaster under dry conditions.     

Soil water content and water relations in planted and naturally regenerated Pinus halepensis Mill. seedlings during the first year in semiarid conditions

The capacity of Aleppo pine (Pinus halpensis Mill.) seedlings to overcome the planting shock in dry conditions was assessed by firstly studying the survival and water status during the first year after planting in relation to soil water content. In spite of receiving only 177 mm rainfall during the year studied, survival of planted Aleppo pine was very high (88.5%). Soil water during summer months (after receiving 67 mm rainfall in winter and spring) was only available at 30–60 cm depth, with tension values of –1.1 and –1.3 MPa in July and August respectively; in these conditions, July predawn xylem water potential measured was –2.5 MPa, and midday potential was –3.6 MPa. According to different authors, these values don't jeopardize the survival of Aleppo pine. In addition, acclimation of outplanted seedlings to environmental conditions was followed by comparing their water relations with those of naturally regenerated seedlings on the site. Predawn and midday xylem water potential showed differences in favour of outplanted seedlings since June, indicating an adjustment to this dry site. Compared to naturally regenerated trees, nursery grown stock of the same age before field planting had much more biomass and higher N and P concentrations and contents; although shoot:root balance and Dickson quality index were not significantly different. Finally, planted seedlings acclimation level during first year was also evaluated by Transplant Stress Index, which value (–0.1278) indicated a slight planting impact.     

Growth, water relations and solute accumulation in osmotically stressed seedlings of the tropical tree Colophospermum mopane

Root and hypocotyl elongation, water status and solute accumulation were studied in osmotically stressed seedlings of the tropical tree, Colophospermum mopane (Kirk ex Benth.) Kirk ex J. Léonard, which grows in hot arid areas of southern and central Africa. Seeds were imbibed for 24 h and then subjected to a polyethylene-glycol-generated osmotic stress of -0.03 (control), -0.2, -0.8, -1.6 or -2.0 MPa for 60 h. Seedlings subjected to moderate water stress (-0.2 MPa) had higher root growth rates (2.41 +/- 0.24 mm h(-1)), greater final root lengths (111 +/- 3.8 mm) and longer cells immediately behind the root elongation zone than control seedlings (1.70 +/- 0.15 mm h(-1) and 93 +/- 3.9 mm, respectively). Root lengths of seedlings in the -0.8 and -1.6 MPa treatments were similar to those of control seedlings, whereas the -2.0 MPa seedlings had significantly shorter roots. Both root and hypocotyl tissues exhibited considerable osmotic adjustment to the external water potential treatments. Seedlings in the -0.03, -0.2, and -0.8 MPa treatments had similar cell turgor pressures (0.69 +/- 0.10, 0.68 +/- 0.07 and 0.57 +/- 0.04 MPa, respectively), whereas the -2.0 MPa treatment lowered cell turgor pressure to 0.17 +/- 0.04 MPa. Root vacuolar osmotic pressures were generally similar to sap osmotic pressures, indicating that the increased root elongation observed in moderately water-stressed seedlings was not caused by increased turgor pressure difference. Neutral-fraction solute concentrations, including the osmoticum pinitol, increased approximately two-fold in root sap in response to a low external water potential. In hypocotyl sap of seedlings in the -2.0 MPa treatment, pinitol more than doubled, sucrose increased from about 2 to 75 mol m(-3) but glucose and fructose remained unchanged and, as a result, total sugars increased only slightly. The benefits of rapid early root elongation and osmoticum accumulation under conditions of water stress are discussed in relation to seedling establishment.     

Water relations and morphological development of bare-root jack pine and white spruce seedlings: seedling establishment on a boreal cut-over site

Bare-root jack pine (Pinus banksiana Lamb.) and white spruce (Picea glauca (Moench) Voss) seedlings were planted on a boreal cut-over site and subsequent growth and seedling water relation patterns were monitored over the first growing season. Comparison of morphological development between white spruce and jack pine showed jack pine seedlings had greater new root development and a lower new shoot/new root ratio, while white spruce seedlings had greater new shoot development. Seasonal water relation patterns showed white spruce seedlings to have a greater decrease in xylem pressure potential (_x) per unit increase in transpirational flux density in comparison to jack pine seedlings. These results suggest that the greater resistance to water flow through the soil-plant-atmosphere continuum in white spruce seedlings compared to jack pine seedlings may be due to the relative lack of new root development in white spruce. Stomatal response of the seedlings showed that as absolute humidity deficit between the needles and air (AHD) increased, needle conductance (gwv) decreased in both species, but at low AHD levels white spruce had gwv approximately 35% higher than jack pine. For white spruce seedlings, gwv decreased as _x became more negative in a predictable curvilinear manner, while gwv of jack pine seedlings responded to _x with a threshold closure phenomenon at approximately - 1.75 MPa. Tissue water potential components for jack pine and white spruce seedlings at the beginning and end of the growing season showed jack pine to reach turgor loss at 76% relative water content while white spruce reached turgor loss at 88% relative water content. White spruce seedlings showed osmotic adjustment over the growing season, with an osmotic potential at turgor loss of - 1.27 MPa and - 1.92 MPa at the beginning and end of the growing season, respectively. Jack pine did not show any osmotic adjustment over the growing season. The implication of morphological development on water relation patterns are discussed with reference to successful seedling establishment.     

Fall lifting and long-term freezer storage of ponderosa pine seedlings: effects on post-storage leaf water potential, stomatal conductance, and root growth potential

Post-storage water relations, stomatal conductance, and root growth potential were examined in ponderosa pine (Pinus ponderosa Dougl. ex Laws.) seedlings from high- and low-elevation seed sources that had been lifted either in October or November and freezer stored, or in March, and then grown hydroponically in a greenhouse for 31 days. Seedlings lifted in October had poor root initiation (< 17 new roots per seedling), low predawn leaf water potentials (< -1.5 MPa), and low stomatal conductance (7.10 mmol m(-2) s(-1)) compared with seedlings lifted in November or March. There was little difference in post-storage water relations and stomatal conductance between seedlings lifted in November and those lifted in March. Throughout the 31-day test, seedlings from the high-elevation seed source produced 3-9 times more new roots, had higher predawn leaf water potentials (-0.6 to -0.7 MPa versus -1.1 to -1.6 MPa), and 1.3-5 times greater stomatal conductance than seedlings from the low-elevation seed source. For all seedlings on Day 31, the number of new roots was significantly related to predawn leaf water potential (r(2) = 0.65) and stomatal conductance (r(2) = 0.82). Similarly, the dry weight of new roots per seedling on Day 31 accounted for a significant amount of the variation in predawn leaf water potential (r(2) = 0.81) and stomatal conductance (r(2) = 0.49).     

Influence of water stress on the physiology and growth of red spruce seedlings

Two-year-old, container-grown red spruce (Picea rubens Sarg.) seedlings from a New Hampshire seed source were exposed to 10 or 11 drying cycles in which the seedlings were not watered until their midday (1400 h) xylem water potentials averaged -1.57 MPa. Control seedlings were kept well watered to maintain midday water potentials of about -0.73 MPa. After the final drying cycle, the water-stressed seedlings were rehydrated and osmotic potentials were determined by pressure-volume analysis. Gas exchange at ambient CO(2) concentration (338 ppm) and at an elevated CO(2) concentration (838 ppm) was measured on both groups of plants as they slowly dried down. No osmotic adjustment or photosynthetic acclimation occurred as a result of the water-stress treatment and both groups of seedlings maintained photosynthesis to water potentials as low as -3.0 MPa. Twenty-four hours after rehydration, the water-stressed seedlings had photosynthetic rates as high as the control seedlings. Estimated stomatal limitation to photosynthesis was approximately 30% down to water potentials of -1.4 MPa, but increased steadily as water potentials decreased further. At ambient CO(2) concentrations (338 ppm) and water potentials averaging -2.45 MPa, photosynthetic rates of water-stressed seedlings were 15% those of well-watered seedlings, whereas when the same water-stressed seedlings were measured in the presence of an elevated concentration of CO(2) (838 ppm) their photosynthetic rates were 73% those of well-watered seedlings measured at an ambient CO(2) concentration (338 ppm).     

Influence of different nursery container media on rooting of Scots pine and silver birch seedlings after transplanting

Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Roth.) seedlings were grown in containers filled with growth media based on medium-textured sphagnum peat, coarse perlite and fine sand. The seedlings were then planted into fine and coarse sandy soils in 2.2 l pots, which were subjected to two water-content treatments (only one for birch). After the seedlings had grown five weeks in a greenhouse, rooting into the surrounding soil and shoot growth were measured. Addition of perlite and sand to peat medium slightly affected rooting; thus suggesting minor effects on seedling establishment. However, nitrogen concentration of the seedlings varied between growth media and correlated positively with rooting into the soil. The particle size and water content of the soil affected considerably rooting of the seedlings. Seedling height at the time of planting did not affect rooting or shoot growth. The fact that the fewest out-grown roots occurred in the dry fine sandy soil, suggests that dry soil together with high strength and resistance to root penetration reduce rooting and water uptake by container seedlings most and may thus cause water and nutrient stresses to seedlings after outplanting.     

Water relations characteristics of competing singleleaf pinyon seedlings and sagebrush nurse plants

Encroachment of singleleaf pinyon (Pinus monophylla) into adjacent low sagebrush (Artemisia arbuscula) and basin big sagebrush (Artemisia tridentata ssp. tridentata) communities may be enhanced by the efficient use of limited water resources by tree seedlings. Seedlings and sagebrush nurse plants were monitored over two growing seasons to determine water-use patterns. Predawn xylem water potential of low sagebrush declined rapidly, reaching −3.5 to −5.5 MPa by late summer. Big sagebrush values dropped to −2.0 to −3.0 MPa during summer drought. The drop in sagebrush xylem water potential was related to the decline in soil water potential (r=0.68 and 0.82). The change in pinyon predawn xylem water potential was moderate, declining to values of −1.5 to −2.5 MPa. An apparent diurnal threshold xylem water potential (−2.3 to −3.0 MPa) that results in stomatal closure enables pinyon seedlings to maintain a seasonally stable xylem water potential. Water use by pinyon seedlings declined by 50% from May to August in association with a reduction in stomatal conductance. Despite reduced stomatal conductance, sagebrush water use continued to increase during summer and reached levels up to five times greater (per unit leaf area) than associated pinyon. Pinyon seedlings appear to have greater drought avoidance than sagebrush nurse plants.     

Loss of hydraulic conductivity due to water stress in intact juveniles of Quercus rubra and Populus deltoides

Despite many studies of the percent loss of hydraulic conductivity in excised branches, there is doubt as to whether cutting stems in air introduces unnatural embolism into the xylem at the cut surface. To address this question, hydraulic conductivity was measured in seedlings of northern red oak (Quercus rubra L.) and rooted scions of eastern cottonwood (Populus deltoides Bartr. ex Marsh.) that had been droughted in pots. Results indicate that in situ dehydration produced a very similar vulnerability curve (% loss of conductivity versus water potential) to those previously obtained by bench-top dehydration of excised branches of eastern cottonwood and red oak. In eastern cottonwood cuttings, conductivity loss increased sharply below water potentials of -1.0 MPa, with 100% loss of conductivity occurring by -2.0 MPa, whereas conductivity loss in red oak seedlings was more gradual, i.e., increasing below -1.5 MPa and sustaining 100% loss of conductivity by about -4.0 MPa.     

Influence of ectomycorrhizal fungal inoculation on growth and root IAA concentrations of transplanted conifers

We determined whether in vitro plant growth regulator production by mycorrhizal fungi is correlated with conifer seedling growth and root IAA concentrations. Container-grown seedlings of interior Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), lodgepole pine (Pinus contorta Dougl.) and ponderosa pine (Pinus ponderosa Dougl.) were inoculated at seeding with ectomycorrhizal fungi having a high, moderate or low capacity to produce either IAA or ethylene in vitro. Inoculated seedlings were grown for one season in the nursery, harvested in December, cold stored over winter and then transplanted to either a nursery field or a forest site in the spring. Seedling morphology and endogenous IAA in roots were measured immediately after cold storage and again six and 12 months after transplanting. Morphological responses to inoculation varied among different mycorrhizal fungi. Free IAA concentration of roots was increased in some inoculation treatments for all conifer species. In seedlings transplanted to a nursery field, in vitro ethylene-producing capacity of the ectomycorrhizal fungi was highly correlated with more morphological features than in vitro IAA-producing capacity. Both IAA- and ethylene-producing capacity were significantly correlated with more morphological features in seedlings transplanted to a forest site than in seedings transplanted to a nursery field. One year after transplanting, only in vitro IAA-producing capacity was correlated with endogenous IAA concentration of roots of the inoculated seedlings. We conclude that growth responses of conifer seedlings can be partially influenced by IAA and ethylene produced by ectomycorrhizal fungal symbionts.     

Canopy development and solar conversion efficiency in Acacia auriculiformis under drought stress

A glasshouse experiment was performed with Acacia auriculiformis seedlings to investigate the effects of decreased soil water potential on phyllode extension, abscission and solar conversion efficiency, e. Six-month-old seedlings were subjected for 39 days to one of four treatments: well watered (soil water potential maintained above -0.5 MPa), moderately drought stressed (soil water potential maintained above -1.5 MPa), severely drought stressed (soil water potential maintained above -2.5 MPa) and well watered but pruned to maintain a leaf area approximating that of the severely drought-stressed treatment. Aboveground biomass accumulation decreased by 21% below that of the well-watered controls in the moderately drought-stressed seedlings and by 47% in the severely drought-stressed seedlings as a result of both decreased interception of solar radiation and lower e. Differences in phyllode extension rate, rather than in phyllode abscission, were primarily responsible for the differences in interception of solar radiation among treatments. Decreases in phyllode extension rate and water use occurred simultaneously in response to decreasing soil water potential. Specific leaf area decreased and water use efficiency increased in response to drought stress.     

Performance of loblolly pine (Pinus taeda L.) seedlings and micropropagated plantlets on an east Texas site: II. Water relations

Loblolly pine (Pinus taeda L.) seedlings and plantlets, produced from drought resistant genetic families were planted in east Texas and their relative water relations compared through the first two, sixth and seventh growing seasons. Stock type (seedling or plantlet) had a larger influence on water relations than did family source in the early years. Seedlings exhibited more of those characteristics generally considered to benefit survival under harsh environments. The principal difference between stock types was the degree of progressive decrease in predawn water potential (Ψ_pre) through the growing season. On average and across families, Ψ_pre decreased through the first growing season by 32 and 17% for plantlets and seedlings, respectively.Complete stomatal closure was never observed for any genetic source or stock type even when water potential was below −2.0 MPa. However, seedlings exhibited more stomatal regulation by lowering their stomatal conductance when water deficit was encountered, and increasing it during periods of favorable soil moisture availability. Predawn water potential was never below −0.9 MPa for any tree in either of the two first growing seasons. Although significant differences between seedling and plantlet performance were observed, their magnitude was small and diminished in the later years (sixth and seventh growing seasons). We conclude that plantlets need further development to be considered as a viable regeneration source for loblolly pine plantations on sites where summer drought is common.     

Pre‐ and post‐inoculation water stress affects Sphaeropsis sapinea canker length in Pinus halepensis seedlings

In order to assess the influence of water stress on the development of Sphaeropsis sapinea cankers in Pinus halepensis, the stems of 4‐ to 5‐year‐old potted seedlings were artificially inoculated with the fungus before and after being kept at controlled water regimes from April 1997 to March 1998. In the pre‐water‐stress inoculation experiment, the canker length, measured 5 months after inoculation (September 1997), was greater in seedlings predisposed to extreme water deficit (midday needle water potential between ?4.5 and ?5.5 MPa). In the post‐water‐stress inoculation experiment, the fungus was inoculated in April 1998, after irrigation had enabled the seedlings to resume normal needle water potential. In this case also, at 5 months after inoculation, longer cankers were visible in seedlings that had been subjected to extreme water deficit. These findings suggest that the occurrence of marked water stress, although apparently tolerated by Aleppo pine, can enhance the development of S. sapinea cankers in this species, regardless of whether the stress occurs before or after infection by the fungus.     

Characterization of BspA, a major boiling-stable, water-stress-responsive protein in aspen (Populus tremula)

We identified a novel 66 kDa boiling-stable protein (BspA) in cultured shoots of aspen (Populus tremula L.) which was highly expressed in response to gradual water stress. The BspA protein, which was highly expressed as early as 1 h after initiation of a drought treatment, accumulated during progressive water stress, decreased on rehydration, and was expressed in response to abscisic acid (ABA) application, as detected by SDS-PAGE protein analysis and Western blotting. Anti-BspA antibodies also cross-reacted with a 119 kDa protein. The 119 kDa protein was also induced by water stress, but it was detected only in the total protein fraction and not in the heat-stable fraction. The BspA protein cross-reacted with antibodies raised against a water-stress-responsive protein isolated from the African resurrection plant Craterostigma plantagineum Hochst. The N-terminal amino acid sequence of BspA was determined and exhibited high homology with the wheat germins GF-2.8 and GF-3.8. The BspA protein was the only major, water-stress-responsive boiling-stable protein detected in aspen.     

Planting stress, water status and non-structural carbohydrate concentrations in Corsican pine seedlings

Two-year-old Corsican pine (Pinus nigra ssp. laricio var. Corsicana) seedlings were either well watered or subjected to a moderate drought for one month before being lifted from the nursery bed on October 9 and transplanted. Well-watered, non-transplanted seedlings served as controls. Needle predawn water potential (Psi(wp)), non-structural carbohydrate concentrations and plant development (survival, bud break, shoot elongation) were assessed before and during the first growing season after transplanting. On April 16, just before bud break, Psi(wp) was lower for the well-watered + transplanted and drought-conditioned + transplanted seedlings (Psi(wp) = -1.45 and -1.83 MPa, respectively) than for the controls (Psi(wp) = -0.56). There was a close relationship between the Psi(wp) measured on April 16 and bud break, shoot elongation and plant survival during the following growing period. Above a Psi(wp) of -1.1 MPa, all plants developed normally. Between -1.1 MPa and -1.6 MPa, bud break, and thus shoot elongation, did not occur in all plants. Between -1.6 MPa and -2.1 MPa, the plants were characterized by the absence of shoot growth, but mortality was zero. Below -2.1 MPa, there was a large increase in plant mortality. On April 16, starch concentrations were markedly lower in the roots of transplanted seedlings than in the controls. There was a positive correlation between Psi(wp) and root starch concentration. The Psi(wp) (-2.3 MPa) at which complete starch depletion was observed in the roots corresponded to the Psi(wp) below which plants did not survive. These results suggest that mechanisms specifically linked to altered water status and metabolic processes associated with altered carbohydrate status are involved in transplanting stress; however, it was not possible to disentangle the two effects. Drought conditioning did not lead to a marked increase in soluble carbohydrate concentrations, as reported for other species, and did not increase plant tolerance to transplanting stress.     

Growth, nutrition and response to water stress of Pinus pinaster inoculated with ten dikaryotic strains of Pisolithus sp

Reconstituted dikaryons of Pisolithus sp. (Pers.) Coker & Couch from South Africa influenced growth parameters (shoot length, shoot/root ratio and leaf area), nutrition and physiological indicators (transpiration rate, stomatal conductance and xylem water potential) of maritime pine (Pinus pinaster Ait.) seedlings during drought and recovery from drought. Seedlings colonized with certain dikaryons were more sensitive to water stress and showed less mycorrhiza formation under water stress than seedlings colonized with other dikaryons. Control (uninoculated) seedlings were significantly smaller than those inoculated with dikaryons. Transpiration rate, stomatal conductance and xylem water potential varied among mycorrhizal treatments during the water stress and recovery periods. After rewatering, the controls and seedlings inoculated with dikaryon 34 x 20 had a weaker recovery in transpiration rate, stomatal conductance and xylem water potential than the other treatments and appeared to have experienced damage due to the water stress. Concentrations of various elements differed in the shoots of Pinus pinaster colonized by the various dikaryons. It is suggested that breeding of ectomycorrhizal fungi could constitute a new tool for improving reforestation success in arid and semi-arid zones.     

Grass interference limits resource availability and reduces growth of juvenile red pine in the field

The effects of competing grasses on resource availability, growth and ecophysiological characteristics of 3-0 red pine (Pinus resinosa Ait.) seedlings were examined the first two years following outplanting in Anoka County, Minnesota, USA. Equal numbers of seedlings were planted into suppressed and undisturbed grass communities in a sandy soil. Grass suppression was maintained throughout the first growing season, but partially discontinued thereafter on the site. During the first field season interference from grass reduced pine seedling root collar diameter, needle length, number of new root tips, and lateral root length by over 40%. Mean pre-dawn needle water potential was 0.55 MPa lower in undisturbed grass plots during a brief drought in year one, but otherwise water stress was not significantly (p=0.05) influenced by grass interference. The presence of grass also reduced, up to 50%, the photosynthetically active radiation reaching the seedling canopy. At the end of year one, total biomass N, P, K, and Ca content were significantly (p=0.05) less in seedlings growing in the undisturbed grass community. Nitrogen was deficient in seedlings growing in grass. After two growing seasons, seedling shoot length (p=0.03), root collar diameter (p=0.001), and needle length (p=0.001) were significantly less (40, 54 and 20%, respectively) for seedlings growing in undisturbed grass. Seedling growth reductions induced by grass competition were associated with multiple environmental stressors in the field and not restricted to water stress as was observed in earlier studies with pine species at low and mid-latitude sites.     

The role of litter and humus layer as a mortality factor of seedlings of Japanese red pine,Pinus densiflora Sieb. et Zucc.

The process of decrease in seedlings of Japanese red pine (Pinus densiflora Sieb.et Zucc.) and the effect of litter and humus layers on the mortality were studied in their stand and adjacent bare land. After the rainy season, only small number of individuals died in the bare land but more seedlings died in the forest. The number of seedlings killed by drought was greater in the forest than in the bare land. The population of naturally occurring current year seedlings was larger in the forest than in the bare land, but that of 1- to 5-year-old seedlings was smaller. Severe summer drought exceeding the permanent wilting point was observed only in the forest F-H horizon where greater part of seedling roots distributed, this may be one of major mortality factors. Inhibitory substances to seedling growth in fresh fallen pine needles were also suggested.     

Potential for Biological Control of Botrytis cinerea in Pinus sylvestris Seedlings

Grey mould (Botrytis cinerea Pers.: Fr.) is the most common economically important fungal disease in Swedish forest nurseries. In tests in a growth room, foliage of predisposed (preinoculation incubation at 35°C for 4 days) Scots pine (Pinus sylvestris L.) seedlings was sprayed with suspensions containing Mycostop®, Binab® TF.WP or GlioMix® at concentrations of 0.5, 1 or 0.5 g l^?1, respectively, and/or conidia of B. cinerea (10⁶ spores ml^?1). Binab and GlioMix reduced grey mould in needles by 94 and 92%, respectively, and were as effective as the fungicide Euparen® M 50 WG, while Mycostop reduced disease by 51%. In one trial in a forest nursery, Mycostop, Binab and GlioMix, each applied two and four times during the growing season, suppressed spontaneous B. cinerea infections in needles of first year container-grown P. sylvestris seedlings by 16–57%, and were as effective as recommended fungicidal sprays. It was concluded that biological control has potential to effectively suppress grey mould in seedlings in forest nurseries.