Effects of Waterlogging on Water Relations of Actively-growing and Dormant Sitka Spruce Seedlings

Two-year-old Sitka spruce [Picea sitchensis (Bong.) Carr.] seedlings,either actively growing or dormant, were waterlogged in a growthroom at 15 °C. Shoot and root growth, transpiration andleaf water potential were observed. In actively-growing plants shoot extension continued after waterlogging,though at a reduced rate, and shoots of dormant plants brokebud and extended during the waterlogging period. Root growthwas suppressed by waterlogging in both types of plant. The 22day waterlogging treatment eventually killed the actively-growingplants but plants which were dormant at the time of waterloggingwere more tolerant. Changes in plant water relations after waterloggingwere entirely different depending on the condition of the plantswhen the soil was flooded. Dormant plants showed a gradual reductionin transpiration and increased water stress over the waterloggingperiod; after the soil was drained leaf water potential increasedto equal the value of control plants which had been maintainedin a freely drained condition, but transpiration did not increaseuntil root growth began. Actively-growing plants exhibited amore complex behaviour, characterized by a very rapid reductionin transpiration after waterlogging, accompanied by a briefperiod of water stress, followed by a period of increasing transpirationrate in the absence of water stress. Finally a second reductionin transpiration occurred and water stress increased as theseedlings died. The importance of the stage of activity of theroot system to the response of plants to waterlogging is discussed. Picea sitchensis (Bong.) Carr., Sitka spruce, waterlogging, water relations, dormancy, transpiration, water potential     

Leaf Water Potential and Control of Water Loss in Droughted Sitka Spruce Seedlings

Sitka spruce seedlings were subjected to drought in experimentsin a growthroom, a greenhouse, and out of doors. The plantswere grown in a double chamber with the bulk of the roots inthe upper part where they dried out the soil when water waswithheld. A few new roots penetrated into the lower part inwhich the soil remained moist. The double chamber system enabledthe plant to attain a high water psotential by night and theshoot was only periodically under mild water stress. Measurementswere made on soil water potential (_solt), leaf water potential(₁), transpiration (E), and stomatal conductance (k_s). As soildecreased over a period of 4.5 d, E and k_s decreased progressively.The decline in E and k_s which indicated stomatal closure, occurredat a higher ₁ than has been reported for Sitka spruce. The behaviourof the stomata appeared to be modified by conditions at theroot, and it is proposed that differences in the response to₁,depend on Whether the latter is reduced by resistances in thexylem between root and leaf, as is known to occur in large treesin moist soil, or by stresses at the root itself.     

Control of Water Loss by Actively Growing Sitka Spruce Seedlings after Transplanting

The roots of actively growing Sitka spruce seedlings were disturbedand damaged to different degrees by transplanting treatmentscarried out in a growthroom. The treatments stopped root extensionand after 2–4 h the transpiration rate of treated plantsdeclined, indicating closure of the stomata, and transpirationcontinued to decrease for a few days. The hydraulic conductivityof the root system was reduced slightly by the treatment butleaf water potential increased or remained unchanged, thereforeno water stress occurred to account for closure of the stomata.Measurements on detached shoots indicated that the water storagecapacity was too small to accommodate changes in the vascularwater supply without reducing leaf water potential. The measuredresponses of the plant were not affected by girdling the baseof the stem prior to root treatment. It is suggested that thetranspiration rate of treated plants was reduced as a resultof a chemical stimulus originating in the root and acting onthe stomata.     

Critical Water Potential for Stomatal Closure in Sitka Spruce

Steady state rates of net photosynthesis and stomatal conductance at high water potentials were measured under controlled conditions in a leaf chamber on Sitka spruce [Picea sitchensis (Bong.) Carr.] shoots detached from the forest canopy or on seedlings. The water supply to the seedlings was terminated by excision and the shoot water potential (or critical water potential) and osmotic potential at the onset of stomatal closure measured. The turgor potential was calculated. The initial osmotic potential before insertion of the shoot into the chamber was also measured. Shoot water potential and osmotic potential at stomatal closure, and initial osmotic potential were significantly higher (less negative) in foliage from the lowest level in the canopy compared with foliage in the upper canopy, and higher in shoots of seedlings transferred to low light than in those at high light. Critical water potential also varied with season, being higher in July than in October and November. In all except one instance, turgor potential at the onset of stomatal closure was negative, possibly because of dilution of the cell sap by the extracellular water during the estimate of osmotic potential. Over all the experiments variation in critical water potential was correlated with variation in critical osmotic potential and, to a lesser extent, the initial osmotic potential. However, turgor potential at the critical potential varied from +0.6 to -4.6 bar. This suggests that difference in turgor between the guard cells and subsidiary cells, which controls stomatal aperture, is only loosely coupled with the bulk leaf turgor and hence that bulk leaf turgor is not a good index of the turbor relations of the guard cells.     

Damage caused by the Green Spruce Aphid to Norway and Sitka spruce needles

Three-month-old needles of Sitka spruce were less susceptible to Elatobium abietinutn than 15-month-old needles. Symptoms appeared after longer aphid feeding times but only a proportion of damaged needles fell. After short feeding periods symptoms appeared in more Norway spruce needles than in Sitka spruce, whereas longer feeding periods resulted in more needles producing symptoms in the Sitka spruce. The symptoms took 4—6 days longer to appear in Norway spruce, and needle fall followed a longer feeding period than on Sitka spruce. Following 72 h feeding, needle fall occurred more quickly on Sitka spruce than on Norway spruce. The time taken for needle fall to occur was inversely related to the feeding time in Sitka spruce but such a response was not evident in Norway spruce. The results are discussed in relation to the differences exhibited in the probing behaviour of the aphid on the two spruces.     

The Effects of Shoot Water Status on Some Photosynthetic Partial Processes in Sitka Spruce

Clonal cuttings of Picea sitchensis (Bong.) Carr. were grown in a controlled environment and, after completion of shoot extension and maturation, subjected to a drying cycle. Photosynthesis and stomatal conductance were measured in situ using ¹⁴CO₂ and a porometer, respectively. Shoot water potential was measured with the pressure chamber. Photosystem and carboxylase activities of chloroplast preparations were measured in vitro. A considerable fall in photosynthetic rate occurred at low water potential. This was associated with stomatal closure and a decrease in CO₂ transfer or fixation processes in the mesophyll. Little change in activity of photosystem I, photosystem II, and ribulose 1,5-diphosphate carboxylase was detected during the drying cycle. Any decline in activity of the photosynthetic partial processes in vivo under severe water stress (Ψ < – 30 bar) was probably masked in vitro as a result of rehydration prior to assay.     

Frost Damage Resulting from Ice Crystal Formation in Seedlings of Spruce and Pine

The influence of ice crystal formation in tissues of unhardened and hardened seedlings of Pinus silvestris L. and Picea abies (L.) Karst. was determined. An apparatus for this purpose was constructed. Unhardened seedlings of spruce collapsed completely as a result of ice crystal formation in their shoots, while unhardened seedlings of pine survived ice crystal formation with or without injury to the needles. After a hardening period of 4–5 weeks, seedlings of spruce and pine survived ice formation but injuries occurred in spruce. For pine, on the contrary, the temperature after ice formation had a decisive effect on the extent of the injuries. Observed injuries are discussed in terms of inter- and intracellular ice crystal formation.     

Leaf Conductance as Related to Xylem Water Potential and Carbon Dioxide Concentration in Sitka Spruce

Current year shoots of Sitka spruce [Picea sitchensis Bong. (Carr.)] from the forest canopy were equilibrated in a leaf chamber. The shoots were excised in air, and removed at differing times in order to establish a relationship between stomatal conductance and xylem water potential. The experiment was repeated at five ambient CO₂ concentrations. A second set of excised forest shoots, and shoots excised from 2-year- old nursery seedlings were allowed to evaporate freely in a controlled environment wind tunnel until a constant rate of transpiration was measured, to establish a relationship between cuticular conductance and xylem water potential. Cuticular conductance was estimated to be 0.012 cm s^-1 at high water potential and declined linearly to 0.007 cm s^-1 at ?3.5 MPa. The implication of this decline in the subsequent calculation of stomatal and mesophyll conductance is considered. Stomatal conductance remained constant at water potentials above ?1.4 MPa and was not affected by ambient carbon dioxide concentrations between 20 and 600 cm^-3. At lower water potentials, stomatal conductance declined and approached zero at ?2.5 to ?2.6 MPa. The results suggest that stomatal aperture is not controlled by either ambient or intercellular space carbon dioxide concentration, and that stomatal closure at low water potential is unlikely to be mediated by carbon dioxide.     

Annual Variations in Competition Symmetry in Even-aged Sitka Spruce

Understanding of the temporal development of competition symmetry,size hierarchy and the role of episodically supplied resourcesin forest stands is limited. Because of intrinsic differencesin the symmetry of competition for light and below-ground resources,it is hypothesized that the overall expression of competitionvaries as a consequence of variations in water availability.Historical growth and precipitation data from an unthinned standof Sitka spruce (Picea sitchensis(Bong.) Carr.) in Denmark areanalysed with respect to the effects of precipitation on competitionsymmetry. The shape and linearity of the size-increment relationshipare analysed, and the temporal development of diameter distributionand size hierarchy is modelled. The test value of lack of linearfit through the origin for the diameter/increment relationshipplotted against precipitation in the growth period forms a triangularenvelope, indicating a more curvilinear relationship for yearswith high water availability. This indicates that dominant treesapparently benefit more from increased precipitation than suppressedtrees—which is also shown in the analysis of the temporaldevelopment of the diameter distribution—and as a consequencethe size hierarchy increases. Finally, it is proposed that temporaland spatial variations in the availability of water and otherbelow-ground resources may have similar effects on competitionsymmetry. Copyright 2001 Annals of Botany Company Competition symmetry, size hierarchy, precipitation, temporal development, Picea sitchensis     

The Effect of Scion Water Potential on Graft Success in Sitka Spruce (Picea sitchensis)

The effect of scion storage on graft success in Sitka sprucehas been examined. Scions kept in cold store for up to 7 d werefound to graft successfully to active rootstocks in about 95%of cases. From 7 d, however, there was a marked decline in successuntil, after 14 d of storage prior to grafting less than 40%of grafts survived. Scion leaf-water potential decreased continuouslyduring storage and fell dramatically during the first 3 d aftergrafting. In successful grafts, scion water potential then recovered,reaching a stable level after 3 weeks. No recovery occurredin grafts, which eventually failed. The relationship betweenscion leaf water potential and graft success rate is discussedwith reference to the role of callus development in graft formation. Picea sitchensis, water potential, graft success     

Effects of Nutrient Heterogeneity and Competition on Root Architecture of Spruce Seedlings: Implications for an Essential Feature of Root Foraging

Background

We have limited understanding of root foraging responses when plants were simultaneously exposed to nutrient heterogeneity and competition, and our goal was to determine whether and how plants integrate information about nutrients and neighbors in root foraging processes.

Methodology/Principal Findings

The experiment was conducted in split-containers, wherein half of the roots of spruce (Picea asperata) seedlings were subjected to intraspecific root competition (the vegetated half), while the other half experienced no competition (the non-vegetated half). Experimental treatments included fertilization in the vegetated half (FV), the non-vegetated half (FNV), and both compartments (F), as well as no fertilization (NF). The root architecture indicators consisted of the number of root tips over the root surface (RTRS), the length percentage of diameter-based fine root subclasses to total fine root (SRLP), and the length percentage of each root order to total fine root (ROLP). The target plants used novel root foraging behaviors under different combinations of neighboring plant and localized fertilization. In addition, the significant increase in the RTRS of 0–0.2 mm fine roots after fertilization of the vegetated half alone and its significant decrease in fertilizer was applied throughout the plant clearly showed that plant root foraging behavior was regulated by local responses coupled with systemic control mechanisms.

Conclusions/Significance

We measured the root foraging ability for woody plants by means of root architecture indicators constructed by the roots possessing essential nutrient uptake ability (i.e., the first three root orders), and provided new evidence that plants integrate multiple forms of environmental information, such as nutrient status and neighboring competitors, in a non-additive manner during the root foraging process. The interplay between the responses of individual root modules (repetitive root units) to localized environmental signals and the systemic control of these responses may well account for the non-additive features of the root foraging process.     

Graft Formation in Sitka Spruce: A Scanning Electron Microscope Study

Low-temperature and conventional scanning electron microscopyhave been used to examine the callus formed at the graft interfacein Picea sitchensis (Bong.) Carr. Callus cells are producedby both cambium and ray parenchyma dedifferentiation and redifferentiationin scion and stock. Adhesion between the cells derived fromscion and rootstock is thought to be my means of pectinaceousbeads at the surface of the callus cells, preceding a more generalfusion of cell walls. The cambia of the two graft componentsare prevented from growing towards each other by the presenceof callus. Instead, the differentiation of new cambium withinthe callus, in the vicinity of the cambia exposed at the preparedsurfaces of the scion and rootstock, links them to form a continuouscambial layer around the combined stem. Callus, cambium, differentiation, grafting, Picea sitchensis     

The Influence of Pine on the Form of Sitka Spruce Fine Roots

The form of fine roots of Sitka spruce (Picea sitchensis Bong.Carr.) when grown in immediate proximity to roots of Scots pine(Pinus sylvestris L.) and when grown separately was comparedusing split-root systems. When spruce roots were intimatelymixed with pine roots the mean length of individual spruce lateralswas significantly greater, while the length: weight ratio andnumber of root tips: weight ratio were smaller than when grownalone. A similar alteration in growth strategy was achievedby direct addition of mineral nitrogen. Key words: Fine root form, pine-spruce interactions     

Dissolved inorganic nitrogen concentrations and fluxes in three British Sitka Spruce plantations

A catchment approach was used to estimate mean dissolved inorganic-N concentrations and fluxes through three mature Sitka spruce plantations at Beddgelert (north Wales), Plynlimon (mid-Wales) and Kershope (Cumbria). Dissolved inorganic-N in bulk precipitation, throughfall, stemflow, soil water at 4 or 5 depths, and streamwater was measured every two weeks for periods of 1 or 2 years at all three sites. Bulk precipitation inputs at the three sites varied from 6 to 10.4 kg N ha^-1 a^-1 and stream outputs varied from 6.4 to 13.6 kg N ha^-1 a^-1. Beddgelert Forest had the highest inputs and outputs and is Nitrogen saturated according to certain Scandinavian criteria (Nilsson, 1986). All three sites had much higher outputs than might be expected from the magnitude of the inputs, since conifer forests are normally regarded as being conservative with respect to N. Only at Plynlimon were reductions in dissolved inorganic N flux or concentration observed from bulk precipitation to throughfall and soil waters. At the other two sites, the tree canopies did not assimilate N in incoming rainfall, and active nitrification resulted in high concentrations of nitrate in soil and streamwater. It is proposed that many mature western upland Sitka spruce plantations may behave in a similar manner with respect to dissolved N, in contrast to sites in eastern Scotland where rates of nitrification are slower and nitrogen appears to be less available to plantation trees.     

Photoactivation of Oxygen-evolving System in Dark-grown Spruce Seedlings

Plastids prepared from dark-grown spruce seedlings showed a negligible activity of photosystem II, and no fluorescence variation was observed during actinic illumination. The photosystem II reaction centre, however, was present in primary thylakoids. Exposure of such seedlings to continuous light induced the development of photosystem II activity via three stages (rapid, lagged and gradual), and the variable fluorescence appeared. The rapid development of photosystem II may be attributed to the activation of the oxygen-evolving system, possibly the manganese-catalyzing site, and the lagged and gradual developments may be closely related to the formation of thylakoid membranes and their assembly to grana.     

Mercuric Chloride Effects on Root Water Transport in Aspen Seedlings

HgCl(2) (0.1 mM) reduced pressure-induced water flux and root hydraulic conductivity in the roots of 1-year-old aspen (Populus tremuloides Michx.) seedlings by about 50%. The inhibition was reversed with 50 mM mercaptoethanol. Mercurial treatment reduced the activation energy of water transport in the roots from 10.82 +/- 0.700 kcal mol(-1) to 6.67 +/- 0.193 kcal mol(-1) when measured over the 4 degrees C to 25 degrees C temperature range. An increase in rhodamine B concentration in the xylem sap of mercury-treated roots suggested a decrease in the symplastic transport of water. However, the apoplastic pathway in both control and mercury-treated roots constituted only a small fraction of the total root water transport. Electrical conductivity and osmotic potentials of the expressed xylem sap suggested that 0.1 mM HgCl(2) and temperature changes over the 4 degrees C to 25 degrees C range did not induce cell membrane leakage. The 0.1 mM HgCl(2) solution applied as a root drench severely reduced stomatal conductance in intact plants, and this reduction was partly reversed by 50 mM mercaptoethanol. In excised shoots, 0.1 mM HgCl(2) did not affect stomatal conductance, suggesting that the signal that triggered stomatal closure originated in the roots. We suggest that mercury-sensitive processes in aspen roots play a significant role in regulating plant water balance by their effects on root hydraulic conductivity.     

Quantum Requirements of Photosynthetic Electron Transport In Sitka Spruce from Different Light Environments

Quantum requirements of photosynthetic electron transport have been measured in shoots of Picea sitchensis (Bong.) Carr. (Sitka spruce) from different levels in a forest canopy and in shoots from plants grown in contrasting light environments in controlled environment chambers. Neutral density filters were used to obtain very low photon flux densities. The light absorbed by the chloroplast suspensions was calculated from measurements of the transmittance of the suspensions. The shoots from the top of the forest canopy (“sun” shoots) had lower quantum requirements for photosystems I and II than the shoots from the bottom of the forest canopy (“shade” shoots). High light grown plants and “sun” shoots had higher rates of electron transport at light saturation than low light grown plants and “shade” shoots. Thus a higher potential for electron transport was found to exist in “sun” shoots than in “shade” shoots at both high and low photon flux densities.     

Genetic diversity in a seed production population vs. natural populations of Sitka Spruce

Isozyme analysis was applied to estimate the level of variation and the genetic structure of a seed-production population (i.e., seed orchard) and 10 range-wide natural populations of Sitka spruce (Picea sitchensis (Bong.) Carr.). Gene diversity and heterozygosity estimates were comparatively high in both the seed orchard and the natural populations studied. The seed orchard population showed a significantly higher number of alleles per locus and percentage of polymorphic loci. Though not significant, mean heterozygosity of the seed orchard was higher than that observed for all natural populations. Genetic distance analysis indicated that the seed-orchard population was genetically similar to three natural populations from which the parent trees were selected. Parent trees sampling breadth has been identified as the major cause for the observed increased level. The impact of recurrent selection and seed orchard biology and management on maintaining the genetic diversity is discussed.