Xylem of early angiosperms: Nuphar (Nymphaeaceae) has novel tracheid microstructure1

SEM studies of xylem of stems of Nuphar reveal a novel feature, not previously reported for any angiosperm. Pit membranes of tracheid end walls are composed of coarse fibrils, densest on the distal (outside surface, facing the pit of an adjacent cell) surface of the pit membrane of a tracheid, thinner, and disposed at various levels on the lumen side of a pit membrane. The fibrils tend to be randomly oriented on the distal face of the pit membrane; the innermost fibrils facing the lumen take the form of longitudinally oriented strands. Where most abundantly present, the fibrils tend to be disposed in a spongiform, three-dimensional pattern. Pores that interconnect tracheids are present within the fibrillar meshwork. Pit membranes on lateral walls of stem tracheids bear variously diminished versions of this pattern. Pits of root tracheids are unlike those of stems in that the lumen side of pit membranes bears a reticulum revealed on the outer surface of the tracheid after most of the thickness of a pit membrane is shaved away by the sectioning process. No fibrillar texturing is visible on the root tracheid pits when they are viewed from the inside of a tracheid. Tracheid end walls of roots do contain pores of various sizes in pit membranes. These root and stem patterns were seen in six species representing the two sections of Nuphar, plus one intersectional hybrid, as well as in one collection of Nymphaea, included for purposes of comparison. Differences between root and stem tracheids with respect to microstructure are consistent in all species studied. Microstructural patterns reported here for stem tracheid pits of Nymphaeaceae are not like those of Chloranthaceae, Illiciaceae, or other basal angiosperms. They are not referable to any of the patterns reported for early vascular plants. The adaptational nature of the pit membrane structure in these tracheids is not apparent; microstructure of pit membranes in basal angiosperms is more diverse than thought prior to study with SEM.     

STIPE ANATOMY,WATER POTENTIALS,AND XYLEM CONDUCTANCES IN SEVEN SPECIES OF FERNS (FILICOPSIDA)

Anatomy and water relations were studied for the desert fern Notholaena parryi, as well as six other ferns representing three different orders which occupied xeric as well as mesic habitats. Tracheid number and diameter, and total xylem cross sectional area increased during leaf development for N. parryi; the whole plant conductance (volume flow of water through a stipe divided by the rhizome-to-leaf water potential drop) increased but tended to level off as the leaves matured. The reported occurrences of very steep water potential gradients (about 25 MPa m^–1) in stipes of N. parryi were confirmed. The ferns with the highest whole plant conductances (Alsophila australis, Botrychium dissectum, and Adiantum capillus-veneris) had the largest or greatest number of tracheids. Numerous tracheids in Botrychium dissectum offset a low tracheary conductivity, whereas Marsilea vestita had few tracheids resulting in a low whole plant conductance. Whole plant conductances for the ferns were 2 to 3 orders of magnitude less than those generally observed for angiosperms and 6 orders less than for gymnosperms. However, the relative conductivity (whole plant conductance times stipe xylem length divided by xylem area) was only 5- to 10-fold less than for angiosperms and about the same as for the gymnosperms. Stipe water relations in these ferns are discussed in relation to the evolution of xylem anatomy.     

Vertical and radial profiles in tracheid characteristics along the trunk of Douglas-fir trees with implications for water transport

The main stems of three young Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirbel) Franco) trees were dissected to obtain samples of secondary xylem from internodes axially along the trunk and radially within each internode. From these samples, measurements were obtained of tracheid diameter, length, the number of inter-tracheid pits per tracheid, and the diameter of the pit membranes. In addition, samples were obtained along the trunks of three old growth trees and also a small sample of roots for measurement of tracheid diameter. A gradient was apparent in all measured anatomical characters vertically along a sequence among the outer growth rings. These gradients arose not because of a gradient vertically along the internodes, but because of the strong gradients present at each internode among growth rings out from the pith. Tracheid characteristics were correlated: wider and longer tracheids had more numerous pits and wider pits, such that total pit area was about 6% of tracheid wall area independent of tracheid size. A stem model combining growth rings in parallel and internodes in series allowed for estimates of whole trunk conductance as a function of tree age. Conductance of the stem (xylem area specific conductivity) declined during the early growth of the trees, but appeared to approach a stable value as the trees aged.     

Morphological adaptations of tracheid structure to water stress gradients in the crown of Sequoiadendron giganteum

Summary Mean radial diameter of tracheids in young branches of a 90 m Sequoiadendron giganteum decreases linearly with height along a gradient correlated linearly with decreasing xylem pressure potential. These smaller tracheid diameters provide strength to resist strong mechanical tensions in the xylem column and hypothetically allow greater efficiencies of water conduction. Tracheid length is not significantly correlated with either water stress or tracheid diameter.     

Biomass and distribution of fine and coarse roots from blue oak (Quercus douglasii) trees in the northern Sierra Nevada foothills of California

This research adds to the limited data on coarse and fine root biomass for blue oak (Quercus douglasii Hook and Arn.), a California deciduous oak species found extensively throughout the interior foothills surrounding the Central Valley. Root systems of six blue oak trees were analyzed using three methods — backhoe excavation, quantitative pits, and soil cores. Coarse root biomass ranged from 7 to 177 kg per tree. Rooting depth for the main root system ranged from 0.5 to 1.5 m, with an average of 70% of excavated root biomass located above 0.5 m. Of the total biomass in excavated central root systems, primary roots (including burls) accounted for 56% and large lateral roots (> 20 mm diameter) accounted for 36%. Data from cores indicated that most biomass outside of the root crown was located in fine roots and that fine root biomass decreased with depth. At surface depths (0–20 cm), small-fine (< 0.5 mm diameter) roots accounted for 71%, large-fine (0.5–2.0 mm) for 25%, and coarse (> 2 mm) for 4% of total root biomass collected with cores. Mean fine root biomass density in the top 50 cm was 0.43 kg m⁻³. Fine root biomass did not change with increasing distance from the trees (up to approximately 5 m). Thus, fine roots were not concentrated under the tree canopies. Our results emphasize the importance of the smallest size class of roots (<0.5 mm), which had both higher N concentration and, in the area outside the central root system, greater biomass than large fine (0.5–2.0 mm) or coarse (> 2.0 mm) roots. This revised version was published online in June 2006 with corrections to the Cover Date.     

Biophysical Model of Xylem Conductance in Tracheids of the Fern Pteris vittata

Calkin, H. W., Gibson, A. C. and Nobel, P. S. 1986. Biophysicalmodel of xylem conductance in tracheids of the fern Pteris vittata.—J.exp. Bot. 37: 1054–1064. Water movement in the xylem is often analysed with the Hagen-Poiseuilleequation, which applies to capillaries of specific diameters.However, the predicted hydraulic conductances per unit length(K_h) are generally much higher than measured values and importantanatomical details, such as the pits of tracheids, are ignored.Here, a previous model based on the Hagen-Poiseuille analysisfor water flow in the stipes of Pteris vittata is improved byincorporating the actual lumen transectional shape (usuallyelliptical or ovate) and the tapering that occurs at the endsof its tracheids, as well as using a better method for analysingthe electrical circuit analogues for the pits (pit cavitiesplus pit membranes). The measured K_h was similar to that predictedby the Hagen-Poiseuille equation for narrow stipes with theirsmall tracheids, but was only about half the measured K^h forlarge stipes. Correcting for the actual shape changed K^h 2-to 3-fold for tracheids with elliptic and ovate transections.For the smaller diameter tracheids, most of the flow resistancewas from the lumens but for the larger tracheids most was fromthe pit membranes. For all stipes the pit cavities accountedfor 12–22% of the total resistance. When the pit membraneswere partially digested away with cellulase, K^h increased about66%, consistent with the deduced resistance of this part ofthe pathway. The present model incorporating realistic anatomicaldetails allowed reasonable predictions of the hydraulic conductanceper unit length over a wide size range of stipes for this fern. Key words: Hydraulic conductance, pit, tracheid, xylem     

WOOD ANATOMY OF BUBBIA (WINTERACEAE), WITH COMMENTS ON ORIGIN OF VESSELS IN DICOTYLEDONS

Quantitative and qualitative features of wood anatomy are reported for ten collections of seven species of Bubbia. Variations on the basic plan for Winteraceae can be interpreted in terms of taxonomic and ecological distinctions. Tracheid length is correlated with plant size and habit: tracheids are shortest in shrubs. Tracheid wall thickness and ray cell wall thickness distinguish species. Ray cell procumbency and multiseriate ray width increase with age. Growth rings occur only in a species from stream margins. SEM studies reveal absence of a warty layer within tracheids. Helical thickenings are absent. Presence of these two features in Pseudowintera may be correlated with the cool temperate habitats of that genus. Overlap areas of tracheids in Bubbia show various degrees of scalariform pitting, ranging from none (B. semecarpoides) to abundant presence (B. balansae). Perforation-like pits in tracheids of the latter prove, with SEM studies, to have pit membranes containing porosities less than 1 μm in diameter. Scalariform pitting on overlap areas is absent in earlier secondary xylem and increases during later secondary xylem. Scalariform lateral wall pitting can occur in abnormally wide tracheids formed after pauses in cambial activity. These facts show that primitive dicotyledon woods like those of Bubbia can activate genetic information for scalariform end wall patterns and lateral wall pitting such as primitive vessels show without the intervention of paedomorphosis. Paedomorphosis in dicotyledon woods is held still to apply only to special herbaceous and herblike growth forms, not to primarily woody plants. Progenesis (in xylem, loss of secondary xylem) is not held to be necessary to account for the scalariform patterns seen in tracheary elements of primitive dicotyledons. Reasons are given for rejection of the hypothesis that Winteraceae and other woody dicotyledons (Amborella, Sarcandra, Tetracentron, Trochodendron) are secondarily vesselless.     

Surface structure and anatomical aspects of Chinese brake fern (Pteris vittata; Pteridaceae)

The objective of this study was to describe a wide spectrum of surface structural and anatomical details of the Chinese brake fern (Pteris vittata) using scanning electron microscopy (SEM). SEM revealed that the epidermal cells of the pinnae were elongated with raised periclinal and sinuous anticlinal walls. The pinnae were hypostomatous with randomly scattered anomocytic stomatal complexes positioned at the same level as the epidermis. Stomates were large and elliptical (27.4 μm × 10.2 μm). Cross sections from the central regions of the rachis and the stipe revealed V- and U-shaped vascular bundles, respectively. In each vascular bundle, the xylem strands were sea-horse shaped (hippocampus). In contrast, the pinnae possessed a triangular vascular bundle with uniform mesophyll organization comprising of homogenous lobed parenchyma cells. The indumentum consisted of trichomes and scales, which formed various types of vestiture. Trichomes were borne only on the pinnae and scales on the rachis and stipe. The roots developed a dense network of long root hairs averaging 244 μm long, and the xylem consisted of tracheids with scalariform pitting. Sori were submarginal; continuous along both margins of the pinna and were covered with a false indusium. The sporangia were oblong with a short thick stalk and the annulus was positioned vertically resulting in transverse dehiscence of the sporangium. The paraphyses were uniseriate, unbranched, septate and found to be intermixed with the sporangia. The exine of the globose spores was adorned with thick reticulum in which the areoles contained round tubercles. This study describes surface features in detail, which is essential to studies examining the issue of whether morphological characteristics are related to arsenic hyperaccumulation inP. vittata.     

Growth and Development of Roots of Grapevine (Vitis vinifera L.) in Relation to Water Uptake from Soil

Developmental patterns of lateral roots and their vascular differentiationwere investigated for Vitis vinifera L. cv. Shiraz to assessthe likely contribution of lateral roots to total water uptakeof plants subjected to different irrigation regimes. Correlationanalyses showed a significant positive correlation between mainroot diameter and the diameter of first order lateral rootsof well-watered plants, but in water-stressed plants the twowere not significantly correlated. The correlations betweendiameters of first order lateral roots and the diameters ofmain roots were greater than correlations between the lengthsof first order laterals and the diameters of main roots. Thesuberised surface area of well-watered main roots increasedfrom 4% of total surface area at 0·25 cm to 100% at 10cm from the tip, whereas that of stressed plants increased from15% at 0·25 cm to 100% at 5 cm from the tip. In all treatmentsthe highest linear density of first order laterals was about7 laterals cm^-1 of main root. More than 50% of first order lateralshad diameters less than 0·05 cm, and more than 90% ofthem had lengths less than 5 cm. Calculations of axial resistancesbased on xylem diameter measurements suggest that the axialresistances of root segments may not be uniform along rootsas is often assumed in models of water uptake. Water flow intothe main roots via the lateral root pathway is likely to bemuch smaller than that via the direct radial flow pathway asonly about 1% of surface area of main roots is directly occupiedby lateral roots, leaving the other 99% of main root surfacearea available for the direct radial flow pathway.Copyright1994, 1999 Academic Press Axial resistance, grapevine (Vitis vinifera L. cv. Shiraz) roots, root diameter, root length, xylem vessels     

Analysis of circular bordered pit function II. Gymnosperm tracheids with torus-margo pit membranes

A model of xylem conduit function was applied to gymnosperm tracheids with torus-margo pit membranes for comparison with angiosperm vessels. Tracheids from 17 gymnosperm tree species with circular bordered pits and air-seed pressures from 0.8 to 11.8 MPa were analyzed. Tracheids were more reinforced against implosion than vessels, consistent with their double function in transport and support. Tracheid pits were 3.3 to 44 times higher in hydraulic conductivity than vessel pits because of greater membrane conductivity of the torus-margo configuration. Tight scaling between torus and pit size maximized pit conductivity. Higher pit conductivity allowed tracheids to be 1.7-3.4 times shorter than vessels and still achieve 95% of their lumen-limited maximum conductivity. Predicted tracheid lengths were consistent with measured lengths. The torus-margo structure is important for maximizing the conductivity of the inherently length-limited tracheid: replacing the torus-margo membrane with a vessel membrane caused stem tracheid conductivity to drop by 41%. Tracheids were no less hydraulically efficient than vessels if they were long enough to reach their lumen-limiting conductivity. However, this may only be possible for lumen diameters below approximately 60-70 μm.     

Dependence of the annual-ring structure in the stem and structural roots of Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) on environmental factors in the forest-steppe zone of Prebaikalia

It was established on the basis of the data of perennial surveys that the annual growth of the layer of early tracheids of the stem and structural roots of pine correlated with each other (r = 0.72−0.73). The sizes of annual rings in the organs studied were determined mainly by the width of the layer of early tracheids (r ² = 83−92%) and were 50% dependent on their radial diameter for the stem and less than 20% dependent for the structural roots. The width of the layer of early and late tracheids in the stem and structural roots was significantly correlated with heat and water availability, and the radial diameter of early and later tracheids was determined mainly by the temperature conditions of the environment. The combined effect of air and soil temperature, air humidity, and precipitation on the annual-ring structure in the stem and structural roots, except on the width of the layer of late tracheids, was 34–58%.     

Very fine roots respond to soil depth: biomass allocation,morphology, and physiology in a broad-leaved temperate forest

Very fine roots (<0.5 mm in diameter) of forest trees may serve as better indicators of root function than the traditional category of <2 mm, but how these roots will exhibit the plasticity of species-specific traits in response to heterogeneous soil nutrients is unknown. Here, we examined the vertical distribution of biomass and morphological and physiological traits of fine roots across three narrow diameter classes (<0.5, 0.5–1.0, and 1.0–2.0 mm) of Quercus serrata and Ilex pedunculosa at five soil depths down to 50 cm in a broad-leaved temperate forest. In both species, biomass and the allocation of very fine roots were higher in the surface soil but lower below 10-cm soil depth compared to values for larger roots (0.5–2.0 mm). When we applied these diameter classes, only very fine roots of Q. serrata exhibited significant changes in specific root length (SRL; m g⁻¹) and root nitrogen (N) concentrations with soil depth, whereas the N concentrations only changed significantly in I. pedunculosa. The SRL and root N concentrations of larger roots in the two species did not significantly differ among soil depths. Thus, very fine roots may exhibit species-specific traits and change their potential for nutrient and water uptake in response to soil depth by plasticity in root biomass, the length, and the N in response to available resources.     

The effects of artificial soil frost on cambial activity and xylem formation in Norway spruce

We studied the effects of artificial soil frost on cambial activity and xylem formation on 47-year-old Norway spruce [Picea abies (L.) Karst.] trees grown on medium fertile site type (with moraine soil) in eastern Finland (62°42′N; 29°45′E). Different soil frost treatments applied were: (1) natural snow accumulation and melting (control, CTRL); (2) artificial removal of snow from soil surface during two consecutive winters (OPEN); and (3) snow clearing and insulation (FROST), which was in other ways similar to OPEN, but the ground was insulated in early spring to delay soil thawing. Each treatment was replicated in three blocks, and two sample trees in each plot were repeatedly microcored during growing seasons of 2006–2007 for the analysis of the onset, cessation and the duration of xylem formation. The phases of tracheid differentiation (tracheids in radial enlargement, secondary cell wall formation, and mature tracheids) were measured from the microcores of 2007. The intra-ring growth and wood density variables were analysed based on X-ray densitometry. In FROST in 2006, xylem formation started a week later than in the other treatments. In 2007, no difference was found between the treatments. The discrepancy in results between the two study years may be explained by between-years variation in weather, i.e., the winter was colder in 2005/2006 than in 2006/2007. No effects of soil frost treatments on tracheid differentiation and on most of the intra-ring growth and density variables were discovered. Our results suggest that the delayed thawing of moraine soil may slightly affect the onset, timing and duration of xylem formation in Norway spruce. However, the effects of delayed soil frost may depend also on the soil type and become more evident with increasing water holding capacity of the soil.     

Root biomass and nutrient dynamics in a scrub-oak ecosystem under the influence of elevated atmospheric CO2

Elevated CO₂ can increase fine root biomass but responses of fine roots to exposure to increased CO₂ over many years are infrequently reported. We investigated the effect of elevated CO₂ on root biomass and N and P pools of a scrub-oak ecosystem on Merritt Island in Florida, USA, after 7 years of CO₂ treatment. Roots were removed from 1-m deep soil cores in 10-cm increments, sorted into different categories (<0.25 mm, 0.25–1 mm, 1–2 mm, 2 mm to 1 cm, >1 cm, dead roots, and organic matter), weighed, and analyzed for N, P and C concentrations. With the exception of surface roots <0.25 mm diameter, there was no effect of elevated CO₂ on root biomass. There was little effect on C, N, or P concentration or content with the exception of dead roots, and <0.25 mm and 1–2 mm diameter live roots at the surface. Thus, fine root mass and element content appear to be relatively insensitive to elevated CO₂. In the top 10 cm of soil, biomass of roots with a diameter of <0.25 mm was depressed by elevated CO₂. Elevated CO₂ tended to decrease the mass and N content of dead roots compared to ambient CO₂. A decreased N concentration of roots <0.25 mm and 1–2 mm in diameter under elevated CO₂ may indicate reduced N supply in the elevated CO₂ treatment. Our study indicated that elevated CO₂ does not increase fine root biomass or the pool of C in fine roots. In fact, elevated CO₂ tends to reduce biomass and C content of the most responsive root fraction (<0.25 mm roots), a finding that may have more general implications for understanding C input into the soil at higher atmospheric CO₂ concentrations.     

Cellular level observation of water loss and the refilling of tracheids in the xylem of Cryptomeria japonica during heartwood formation

The mechanism of heartwood formation in Cryptomeria japonica D. Don has long been studied since heartwood formation is a fundamental physiological feature of trees. In this study, the water distribution in the xylem of C. japonica was investigated at the cellular level to reveal the role of water distribution in the xylem during heartwood formation. Samples were taken from different heights of each trunk, in which the phases of heartwood formation differed. These were designated as SIH, which consisted of sapwood, intermediate wood, and heartwood; SI, which consisted of sapwood and intermediate wood but no heartwood; and S-all, which consisted entirely of sapwood. Cryo-scanning electron microscopic observations of the heartwood-formed (SIH) and non-heartwood-formed (SI and S-all) xylem revealed different patterns of water distribution changes in tracheids between the latewood and earlywood. In the latewood, almost all tracheids were filled with water in all areas from the sapwood to the heartwood (98–100% of tracheids had water in their lumina). In the earlywood, however, the water distribution differed between the sapwood (95–99%), intermediate wood (7–12%), and heartwood (4–100%). Many of the tracheids in the xylem, where the sapwood changed to intermediate wood lost water. In the heartwood, some tracheids remained empty, while others were refilled with water. These results suggest that the water distribution changes in individual tracheids are closely related to heartwood formation. Water loss from tracheids may be an important factor inducing heartwood formation in the xylem of C. japonica.     

Extracellular Matrix in Early Stages of Direct Somatic Embryogenesis in Leaves of Drosera spathulata

The growth and water relations of Paulownia fortunei in photoautotrophic cultures (nutrient medium lacking sucrose and growth regulator) with CO₂ enrichment (PWAH) or without CO₂ enrichment (PWAL) were compared with those in photomixotrophic shoot (PWC; 30 g dm⁻³ sucrose and 0.3 mg dm⁻³ N⁶-benzyladenine) and root cultures (PWR; 0.3 mg dm⁻³ indole-3-butyric acid). The photoautotrophic and photomixotrophic cultures were incubated under photosynthetic photon flux 125 and 60 μmol m⁻² s⁻¹, respectively. 100 % sprouting and significantly higher number of shoots (1.6) were obtained with PWAH as compared to PWAL and PWC. PWAH and PWAL stimulated spontaneous rooting from the cut end of axillary shoots. In PWAH, 84 % of shoots rooted with an average of 5.9 roots per shoot and 4.0 cm of root length in 21 d. Rooting of photomixotrophic shoot cultures were stimulated by an auxin treatment. In this case, 98.3 % of shoots were rooted with an average of 4.6 roots per shoot and 1.9 cm length. A microscopic observation on leaf abaxial surface prints from photomixotrophic shoot and root cultures showed widely open (6 – 8 μm) spherical stomata (12 – 14 μm) and from photoautotrophic cultures elliptical stomata (10 – 12 μm) with narrow openings (3 – 4 μm). Leaves from photomixo-trophic cultures had higher stomatal index as compared to photoautotrophic cultures. The rate of moisture loss from detached leaves was not varying significantly in different cultures. This revised version was published online in July 2006 with corrections to the Cover Date.     

Immunohistochemical, Histochemical and Radioassay Analysis of Nitric Oxide Synthase Immunoreactivity in the Lumbar and Sacral Dorsal Root Ganglia of the Dog

Summary In this study, immunohistochemistry for neuronal nitric oxide synthase (bNOS-IR), nicotinamide adenine dinucleotide phosphate diaphorase histochemistry (NADPHd) and nitric oxide synthase radioassay were used to study the occurrence, number and distribution pattern of nitric oxide synthesizing neurons in the lumbar (L1–L7) and sacral (S1–S3) dorsal root ganglia of the dog. Nitric oxide synthase immunolabelling was present in a large number of small- (area <1000 μm²) and medium-sized (area 1000–2000 μm²) as well as in a limited number of large-sized (area >2000 μm²) neurons. Although neuronal nitric oxide synthase immunolabelling and histochemical staining provided intense staining of multiple small- and medium-sized neurons in all lumbar and sacral dorsal root ganglia, immunolabelled or histochemically stained somata exhibited little topographic distribution in individual dorsal root ganglia. Great heterogeneity was noticed in the immunolabelling of medium-sized nitric oxide synthase immunopositive neurons ranging from lightly immunolabelled somata to heavily immunoreactive ones with completely obscured nuclei. Both staining procedures proved to be highly effective in visualizing intraganglionic fibers of various diameters. In general, the largest fibers revealed at the peripheral end of lumbar and sacral dorsal root ganglia were larger, 6.49–9.35 μm in diameter, while those running centrally and proceeding into the dorsal roots were about 30% reduced, ranging between 5.32 and 8.67 μm in diameter. Peripherally, the occurrence of nitric oxide synthase detected in axonal profiles, and confirmed histochemically, in the specimens of the femoral and sciatic nerves, is the first indication of the presence of nitric oxide synthase in the peripheral processes of somata located in L4–S2 dorsal root ganglia. Large and thin central nitric oxide synthase immunoreactive processes of L1–S3 dorsal root ganglion neurons segregate shortly before entering the spinal cord, the former making a massive medial bundle in the dorsal root accompanied by a slim lateral bundle penetrating Lissauer's tract. Quantitative assessment of the distribution of bNOS-IR and/or NADPHd-stained neurons showed a peculiar pattern in relation to spinal levels. Apparent incongruity was found in the total number of NADPHd-stained versus bNOS-IR neurons, demonstrating a clear prevalence of small bNOS-IR somata in all lumbar ganglia, while medium-sized NADPHd-stained somata clearly prevailed all along the rostrocaudal axis with a peak in L5 ganglion. While the number of small bNOS-IR neurons clearly outnumbered NADPHd-stained and NADPHd-unstained somata in S1–S3 ganglia, an inverse relation appeared comparing the total number of medium-sized NADPHd-stained and NADPHd-unstained somata compared with the number of moderate and intense bNOS-IR neurons. Densitometry of bNOS-IR and NADPHd-stained neurons in lumbar and sacral ganglia revealed two distinct subsets of densitometric profiles, one relating to more often found medium-sized bNOS immunolabelled and the other, characteristic for moderately bNOS immunoreactive somata of the same cell size. Considerable differences in catalytic nitric oxide synthase activity, determined by conversion of [³H]arginine to [³H]citrulline were obtained in lumbosacral dorsal root ganglia all along the lumbosacral intumescence, the lowest (0.898± 0.2 dpm/min/μg protein) being in the L4 dorsal root ganglion and the highest (4.194± 0.2 dpm/min/μg protein) in the S2 dorsal root ganglion.     

Ammonification in soil aggregates: Effect of some physical,physico-chemical and chemical properties

Aggregate diameter affected significantly the intensity of ammonification in chernozemic rendzina but not in lessivē soil. In the latter the process was influenced significantly by the number of microorganisms able to grow on asparagine agar. A high correlation, though not significant at the level of 0.05, was found between the ammonification intensity and the content of pores of radius: 3–1.5, 7.5–5.0, 0.5–0.25 and 0.01–0.005 μm in chernozemic rendzina and those measuring 1.5–0.5, 0.025–0.01, 0.01–0.005 and >7.5 μm in lessivē soil aggregates as well as the percentage of soil particles of 100–50 μm in chernozemic rendzina aggregates and the internal surface area and organic C in aggregates of lessivē soil.     

Distribution,length and weight of roots in young plantations ofEucalyptus grandis W. Hill ex Maiden irrigated with recycled water

Summary The root systems ofEucalyptus grandis W. Hill ex Maiden, irrigated with recycled municipal effluent at two sites in north-western Victoria, Australia, were studied by excavation and coring. Trees at Robinvale were four years-old and were irrigated using micro-sprays that covered only 70% of the ground surface area, whereas at Mildura, effuent was uniformly was uniformly applied to six years-old trees by flood and sprinkler irrigation. At Mildura where roots were excavated from a 2.80×2.80×1.20 m block of soil, a total root length of 1193 m.m⁻² and a total root weight of 3.1 kg m⁻² were estimated in the top metre. For roots >1 mm diameter, 77% of intercepts were at 0–30 cm, whereas only 50% were in the 50–100 cm soil horizon. At both sites where roots in the top 30 cm were studied by coring, the vertical distributions of root intercepts, length and weight were similar. Root length was greatest in the 0–10 cm soil horizon at both sites, and intercepts of roots <1 mm diameter comprised 73% and 81% of all roots at Mildura and Robinvale respectively. Roots <1 mm diameter contributed 85% of total length at both sites, but only 19% and 21% of total weight at Mildura and Robinvale respectively. The horizontal distribution of roots differed at the two sites. With uniform application of effuent at Mildura, root intercepts and length were concentrated in the centre of the irrigation bay, but at Robinvale, the concentration occurred closer to the tree row due mainly to the different method of irrigation. Root weight at both sites was highest within 50 cm of the tree row. Root densities of 0.11 to 0.57 cm cm⁻³ were estimated in the two plantations; these were similar to root densities measured inPinus radiata D. Don plantations up to 46 months old, but were considerably lower than those estimated for pastures. The implications of the results for the management of irrigated plantations of eucalypts are discussed.