A walk through the forest is a good way to become aquainted with the variety of terpenes emitted by plants. The most fragrant of these are the monoterpenes, which lend a distinctive smell to trees (e.g. α- and β-pinene from pines), mints (e.g. menthol from peppermint), fruits (e.g. limonene from citrus) and flowers (e.g. geraniol from roses). In addition to adding fragrance, phytogenic hydrocarbons play an important role in atmospheric chemistry. This was recognized as early as 1960 by Fritz Went who noted that blue hazes often formed above forested areas (Went, 1960). This idea was picked up by US President Ronald Reagan who accused trees of polluting the atmosphere, although in truth phytogenic hydrocarbons only contribute to pollution in air that is already contaminated by industry or automobile exhaust (Feshenfeld et al., 1992).

Terpenes serve a variety of functions in plants including deterring herbivores and attracting pollinators. Until recently, the function of terpene emission from some groups of plants, which do not store the terpenes in their tissues, has been a mystery. Following the discovery that isoprene, a hemiterpene, increases the thermotolerance of photosynthesis in some species (Sharkey & Singsaas, 1995; Singsaas et al., 1997), Loreto et al. (1998) discovered that thermotolerance of the monoterpene-emitting oak, Quercus ilex, increases when leaves are fumigated with monoterpenes. Now, a report by Delfine et al. in this issue (pp. 27–36) provides further support for the hypothesis that terpenes play a role in thermotolerance of leaves and suggests, intriguingly, that the mechanism by which this occurs is not limited to plants that make these compounds.

For wood mycologists, the work of Savory (1954a,b) marked an important step in the understanding of decay processes by lignolytic fungi. His description of decay by ascomycete and deuteromycete fungi in wood from industrial water cooling towers revealed a particular pattern observed previously by Bailey & Vestal (1937), Tamblyn (1937) and Barghoorn & Linder (1944) in wood from other situations. This type of decay was termed ‘soft rot’ because of the spongy texture of the wood surface. Soft rot, characterized by hyphal penetration of the wood cell walls in a fashion similar to sap-stain fungi, results in the formation of decay cavities in the central zone of wood cell walls, which leads to their ultimate destruction. Soft rot was therefore described as distinct from the brown and white rot forms of wood decay normally attributed to lignolytic basidiomycetes and some of the larger ascomycetes, such as members of the Xylariaceae. Indeed, members of the genera Daldinia, Hypoxylon and Xylaria have usually been considered to be white rot fungi. However, the boundaries between these three types of fungal decay have since become less clear-cut and a report in this issue, of a study into hyphal penetration of the reaction zone in beechwood (Schwarze and Baum, pp. 129–140), identifies different penetration mechanisms.

Of the three component organs of the grass leaf – blade, sheath and ligule – the ligule is the least studied and the least understood. Traditionally, it has been assumed to act in a passive way in protecting the culm and leaves that it encloses from the entry of water, dust and harmful spores. However, ultrastructural and cytochemical studies of the membranous ligules of several taxa, particularly Lolium species, have challenged that rather simplistic view and suggest that these ligules play a more active role in the life of the grass plant as a secretory tissue. This review summarizes the evidence for the latter notion, assesses the validity of both the passive and active hypotheses of membranous ligule function and notes similarities between membranous grass ligules, root caps and lycopsid ligules.

We tested if fumigation with exogenous monoterpenes might induce thermotolerance in leaves of an oak species (Quercus suber) which does not form and emit isoprenoids. To understand if exogenous monoterpene fumigation results in internal accumulation of monoterpenes, a physical method of monoterpene extraction was used. The internal content of monoterpenes increased in concert with increasing fumigation doses. This unambiguously demonstrated acquisition of exogenous monoterpenes. We exposed fumigated Q. suber leaves to two cycles of increasing temperatures from 35 to 55°C at 5°C steps. When leaves were exposed to a low dose of exogenous monoterpenes, yielding an internal content similar to that endogenously formed in the leaves of the monoterpene-emitter Q. ilex, no clear improvement in thermotolerance was found. When leaves were exposed to a high dose of exogenous monoterpenes, yielding an internal content of about five fold the endogenous pool of Q. ilex, but comparable with the expected content following stress-induced stomatal closure, photosynthesis inhibition at high temperatures was attenuated. This effect was observed only at temperatures <45°C during the first cycle, but at all temperatures between 35 and 55°C when plants were exposed to two cycles of high temperatures. Monoterpenes were still found in the leaves of Q. suber 12 h after ending the fumigation. Monoterpenes were also found in non-fumigated leaves distant up to 45 cm from the fumigated leaves. If monoterpenes make the photosynthetic apparatus more resistant to high temperatures, the effect might not be limited to the fumigated leaves and might be persistent after fumigation.

We conducted two experiments to investigate the expression of shade avoidance in response to low ratios of red to far-red irradiation (R[ratio ]FR) in the C4 perennial grass Schizachyrium scoparium at two levels of above-ground (photosynthetic photon flux density (PFD) 400–700 nm) and below-ground (soil volume) resource availability and in plants of two ages. Young plants showed greater sheath and ramet height in response to low R[ratio ]FR and old plants showed reduced ramet initiation in experiments one and two, respectively, but both responses were not expressed simultaneously in either group. Growth of all ramet variables, including ramet initiation, was suppressed in small as opposed to large soil volumes. By contrast, architectural variables, but not ramet initiation, were greater for plants grown in low as opposed to ambient PFD. However, expression of shade avoidance to low R[ratio ]FR was not significantly affected by either level of PFD or soil volume. We must conclude that the levels of resource availability provided do not modulate shade avoidance in this perennial grass. These results demonstrate that S. scoparium is capable of expressing shade avoidance in response to low R[ratio ]FR, but inconsistent juvenile ramet initiation and architectural responses in plants of different ages and phenological stages of development indicate that the shade avoidance response might not be expressed consistently throughout the life of plants.

This paper reports the effects of nutrient magnesium (Mg) concentrations on the growth and photosynthetic physiology of clonal Pinus radiata from four female parents (families) known to differ in their tolerance to Mg deficiency and in their needle Mg concentrations. Plants were grown in flowing nutrient solutions with 2 mg l−1 (control) and 0.8 mg l−1 (low) Mg. Plant growth, needle Mg concentration, photosynthesis, chlorophyll fluorescence and carotenoid pigment content were measured. At low Mg, needle Mg concentration was about half that of control plants, height growth was reduced 15–25%, and the needles showed strong visual characteristics of Mg deficiency. Photosynthesis was also halved, and was associated with closure of the stomata under low Mg and with reductions in the residual conductance. In needles from plants grown at low Mg, photochemical yield was reduced both in the light and in the dark, and was strongly dependent on needle Mg concentrations below a threshold concentration of 0.02–0.025% (d. wt basis). The electron transport rate (ETR) at saturating photon flux density in low-Mg-grown needles was reduced to about half that of their Mg controls, but the photon efficiency of ETR was unaffected by the Mg concentration the plants were grown in. Photosynthetic quenching was markedly reduced and non-photosynthetic quenching was increased following growth in low Mg. Growth under low Mg also increased levels of zeaxanthin. Although family differences in growth and photosynthetic physiology were present, few family × Mg interactions were significant. We conclude that Mg deficiency probably affects growth through severe reductions in photosynthesis.

Ring widths of five Mediterranean forest tree species (Arbutus unedo, Fraxinus ornus, Quercus cerris, Quercus ilex and Quercus pubescens) growing close to a natural source of CO2 in Tuscany, Italy and at a nearby control site were compared. At the CO2-enriched site, trees have been growing for decades under elevated CO2 concentrations. They originated from parent trees that also grew under elevated CO2 in natural conditions, and they have been continuously exposed to elevated CO2 throughout their growth. Tree-ring series from each of the species were prepared. Assigning calendar dates to rings was difficult but possible, and ring-width series were built for all species. The ring-width data were analysed using a two-sided t-test to assess if there was a difference between the radial growth at the CO2-enriched site and the control site. The cumulative basal area at the same cambial age at both sites was also compared using a Wilcoxon test. Radial growth of trees at the CO2-enriched site was not significantly different from growth at the control site. For each species, year by year, radial growth at the CO2-enriched site was tested against the control site and significant differences were found in only a few years; these differences were not synchronous with extreme climatic events. The expected increase in above-ground productivity, as one of the ecosystem responses to increasing CO2 during drought stress, was not observed in this Mediterranean woody plant community, despite being water-limited. Other resource limitations, such as low nutrient availability (common in the Mediterranean region), may have counteracted the positive effect of elevated CO2 under drought stress, or trees may have acclimated to the high CO2.

Plants of Molinia caerulea were grown in sand culture for two seasons. All nitrogen (N) supplied to the plants was enriched with 15N throughout the first season, and at natural isotopic abundance throughout the second season. A series of destructive harvests was taken during the second season. At each harvest, the N mobilized from roots and swollen basal internodes and the protease activity (ability to degrade azocasein) of these storage tissues were measured. The pH response curves of protease activity of both basal internodes and roots exhibited optima at pH 5 throughout the season. The protease activity of roots and basal internodes increased in spring, both on a unit fresh weight and unit protein basis, concomitant with mobilization of N from both these tissues to new shoot growth. In absolute terms more N was mobilized from roots than from basal internodes. However, basal internodes which, compared with roots, showed the greatest protease activity (on a f. wt or unit protein basis) were also the tissues that mobilized a greater proportion of N present in the tissue over winter to new shoot material. Individual clones of M. caerulea varied both in the amount of N mobilized from roots and in root protease activity (per plant). Individuals with a higher protease activity mobilized more N compared with individuals of lower protease activity. Therefore in M. caerulea, relationships between N mobilization and protease activity exist at several levels: (1) between different tissues; (2) temporally throughout the season; and (3) between individual clones.

Roots of angiosperms differ in the behaviour of their apical meristems, and this diversity has been studied through the patterns of differentiation in a range of families. Except for three families of the Nymphaeales, all dicotyledons derive the epidermis wholly or partially with the root cap. In closed meristems the inner cell layer of the cap complex forms the epidermis; in open meristems, where cortex and cap are intermittently of common origin, linkages between the epidermis and the cap predominate. Roots of most monocotyledons derive the epidermis as the outer layer of the cortex. Nymphaeaceae, Cabombaceae and Barclayaceae resemble monocotyledons in this respect, and this applies even in the species with open meristems, in which a cleft forms between the cap and the rest of the apex. This feature fits with the close relationship between the Nymphaeales and monocotyledons revealed by other data. But none of the other Magnoliidae with close links to Liliidae has this type of epidermis. The Nymphaeales are also almost unique among dicotyledons in having trichoblasts in a vertical pattern. This results from transverse divisions of epidermal cells giving two conspicuously different daughters. The smaller and most densely staining one later bears a root hair. This is the way trichoblasts develop in many monocotyledons. The only other dicotyledons found with trichoblasts in vertical patterns are in the Piperaceae, though their closed meristems derive the epidermis in the normal dicotyledon pattern. All other dicotyledons that bear trichoblasts develop them in radial patterns probably related to the radial disposition of cells across the cortex. This pattern is more widespread than was previously known, though it can occur only in roots with closed meristems. In plants of both subclasses with vertical patterns of trichoblasts, it is usually the proximal daughter of the initiating mitosis which becomes the trichoblast. But it is the distal daughter in Juncaceae, Restionaceae, Cyperaceae and Poaceae. Vertical patterns differ further as to whether the atrichoblast daughters divide after initiation, thus changing the alternation of trichoblasts and atrichoblasts along a file of epidermal cells. Monocotyledons and the Nymphaeaceae can produce trichoblasts in open or closed meristems. In some embryos and root primordia the epidermis becomes discrete from cortex and cap and remains so in the meristems. This has now been found in six aquatic genera of the Liliidae, but reports of it occurring elsewhere are probably due to misinterpretation of sections. Discrete epidermises differentiate into trichoblasts and atrichoblasts in Hydrocharitaceae, but not in Araceae or Lemnaceae.

Parameters of convective ventilation and also the content and composition of free amino acids and free carbohydrates in roots, basal culm internodes and leaves were investigated in five stands of Phragmites australis that were subjected to different growth conditions and salinity levels. Shoot length, stem diameter and productivity decreased with increasing salinity. These morphometric variations in turn affected parameters of convective ventilation and amino acid patterns. Phragmites populations growing at the highest salinity levels possessed the lowest rhizome conductivities, which was probably caused by their more pronounced xeromorphic growth. High counterpressures of the rhizome system and low stem diameters were limiting factors for convective flows. The Phragmites stand with lowest flow rate and ventilation efficiency had the highest percentage of alanine, serine and γ-aminobutyric acid (55% of total free amino acids) in its roots, indicating hypoxic stress in basal plant parts. Increasing salinity was connected with significantly higher fractions of glutamine (in basal culm internodes), glutamate (in roots) and proline (in both) as well as slightly increasing osmolality of plant sap. However, total contents of free amino acids and carbohydrates were low, and not related to either salinity level or osmolality of plant sap. This suggests that they might not have an important, direct role as an osmolyte in P. australis. Stand-specific variations in amino acid and carbohydrate patterns of leaves were negligible. Ratios of amino acids to carbohydrate were positively related to potential N supply. It is concluded that (1) salinity affects convective rhizome ventilation by varying the reed morphometry, and (2) the pattern of free amino acids responds characteristically to conditions of salinity and hypoxic growth.

Companion cells in the secondary phloem of 125 Indian dicotyledonous species belonging to 43 families were examined by light microscopy. Four types of companion cell were identified: E-, S- and L-types which were equal in length, shorter, or longer, respectively, than the associated sieve tube element; and R-type, in which two or more companion cells in a vertical row were associated with the sieve tube element. The commonest was the E-type and the rarest was the L-type. E-type companion cells were most frequently found associated with short sieve tube elements (50–250 μm) which had a high frequency of simple sieve plates, considered phylogenetically advanced. R-type companion cells were most frequently associated with long sieve tube elements (>400 μm) with a high frequency of compound sieve plates, considered phylogenetically the least advanced. A strong positive correlation was found between the average number of companion cells associated with a sieve tube element and the lengths of the sieve tube elements. There was also a strong negative correlation between the average number of companion cells associated with a sieve tube element and companion cell lengths.

Within the growth zone, salt-affected leaves of sorghum (Sorghum bicolor) had narrower protoxylem and metaxylem cells than controls. Leaf width and cross-sectional area were also reduced, so that the salt treatment had no effect on the area of protoxylem per area of leaf cross section. Dye uptake studies suggested that in controls most of the veins, but in salt-affected leaves only half of the veins, are functional in water transport. Volumetric water flow was greatly diminished in the salt-affected plants. The reduced flow rate was largely explained by the salt-induced decrease in leaf surface area. Some decreases in flow rates per unit leaf mass or area were also produced by salinity, particularly in late developmental stages. Not surprisingly, leaf conductance measured with a diffusion porometer did not appear to be correlated with the diameter of the protoxylem or metaxylem elements. By contrast, published values of water deposition rates are strongly related to the size of the protoxylem elements: the rates of water deposition into the growing leaf tissue are proportional to the square of the protoxylem radius. Thus environmentally produced change of the hydraulic architecture of monocot leaves may cause change in local growth rates.

Fungal growth within reaction zones of beech (Fagus sylvatica) challenged by three basidiomycetes, Inonotus hispidus, Ganoderma adspersum, Fomitopsis pinicola, and one ascomycete, Ustulina deusta, was studied in naturally colonized and artificially inoculated wood. All the fungi, except F. pinicola, breached reaction zones, but the mechanisms involved were all somewhat different. Both I. hispidus and U. deusta bypassed blocked cell lumina by tunnelling through cell walls (soft-rot mode), but the latter caused far more decomposition of cell walls. Degradation of polyphenols was slight with I. hispidus and absent with U. deusta. By contrast, G. adspersum preferentially degraded the polyphenolic occlusions in the cell lumina. The failure of F. pinicola to invade reaction zones was typical of a brown rot fungus having limited enzymatic potential and a uniform growth pattern. Mechanisms of lesion expansion, illustrated and summarized in schematic diagrams, are consistent with earlier observations that reaction zones in beech sapwood are static boundaries, which may be successfully breached by white and soft-rot fungi.

A new stripe rust resistance gene, YrH52, derived from the unique Mount Hermon population of wild emmer wheat (Triticum dicoccoides) in Israel, was previously located on chromosome 1B. The main objectives of the present study were to increase marker density in the vicinity of the YrH52 gene using additional microsatellite markers, and to evaluate the accuracy and efficiency of marker-assisted selection on this gene in the F2 generation. By means of 70 additional microsatellite primer pairs, 150 individuals of the F2 mapping population were genotyped. Among 202 marker loci, 20 were found to be linked to the YrH52 gene with log-likelihood (LOD) scores ranging from 3.84 to 58.82, and linkage distances ranging from 0.33 to 41.39 cM. A genetic map was constructed of chromosome 1B, consisting of 23 markers and the YrH52 gene, with a total map length of 149.5 cM. Most of the markers were located in the region close to YrH52, which was flanked by Xgwm413 and Xgwm273a with map distances of 1.3 and 2.7 cM, respectively. The accuracy and efficiency of marker-assisted selection were calculated as AMAS and EMAS, respectively, for homozygous resistant genotypes of YrH52 gene in the F2 generation. AMAS and EMAS for homozygous resistant genotypes of the YrH52 gene in the F2 generation showed linear and significant negative correlation with the map distance between marker and target gene or between the two bracketing markers. It was concluded that the YrH52 region on chromosome 1B is significantly more enriched by microsatellite markers than the previously published map; that a single microsatellite marker is efficient for marker-assisted selection of homozygous resistant genotypes of YrH52 gene in the F2 generation when the map distance is <5.0 cM; and that when two markers are used AMAS can be dramatically improved and becomes relatively stable, whereas EMAS will not be obviously improved and will still vary linearly with map distance.

We studied the transport of 15N from a soil compartment separated from a plant root compartment by a hydrophobic polytetrafluoroethylene (PTFE) membrane to plants in the presence and absence of arbuscular mycorrhizal fungi (AMF). We have previously shown that this type of membrane efficiently inhibits mass flow and diffusion of mobile ions in the soil solution in an abiotic system, but can be penetrated by the hyphae of mycorrhizal fungi. Mycorrhizal tomatoes (Lycopersicon esculentum) colonized by Glomus mosseae were grown at two N fertilizer concentrations in a root compartment. A PTFE membrane was placed between the root compartment and an adjoining soil compartment that was inaccessible to the roots but accessible to the AMF hyphae (hyphal compartment). Additional N was applied to the hyphal compartment using uniformly 15N-labelled NH4NO3. There was a flux of 15N from the hyphal compartment to the plants even in the absence of mycorrhizal fungi. However, this flux was much higher in mycorrhizal plants, which had much higher N concentrations in their shoots and roots than did the non-mycorrhizal control plants. This was particularly apparent when the root compartment had a low N fertilizer concentration. Of the total N content of mycorrhizal plants, c. 42 and 24% at the low and high N fertilizer concentrations, respectively, were estimated to originate from the hyphal compartment by transport through AMF hyphae. In the presence of mycorrhizal fungi, the flux of 15N was about three times higher than in their absence. The results show that AMF can access a soil compartment separated by a PTFE membrane, and can contribute substantially to N uptake by plants.

Ectomycorrhizal seedlings of Scots pine (Pinus sylvestris) inoculated with the nitrotolerant Laccaria bicolor and the nitrophobic Suillus bovinus were exposed to ambient (350 μl l−1) and elevated (700 μl l−1) [CO2]. After 79 d the seedlings were labelled for 28 d with 14CO2, after which they were harvested. 14C was determined in shoots, roots plus mycorrhizas, soil, and below-ground respiration; nitrogen was determined in shoots and roots. Total net 14C uptake increased under elevated [CO2]. The extra carbon did not increase the shoot mass but was translocated to the roots and resulted in a decreased shoot-to-root ratio in the Suillus-inoculated seedlings. Laccaria-inoculated seedlings did not incorporate the additional carbon in root or fungal tissue but only increased below-ground respiration. S. bovinus acquired or transferred nitrogen better than L. bicolor and enabled the seedlings to perform better with regard to net carbon uptake under elevated [CO2]. This resulted in nitrogen concentrations in shoots of Suillus-inoculated seedlings that were twice as high as in Laccaria-inoculated seedlings, irrespective of [CO2]. The higher nitrogen concentration in the shoots resulted in a doubling of the 14C uptake per unit shoot mass. Our results suggest that the ability of ectomycorrhizal Scots pine seedlings to respond positively to elevated atmospheric [CO2] is strongly fungal-species specific.

The non-reducing disaccharide trehalose is a major storage compound in most fungi. For a better understanding of carbon metabolism in the ectomycorrhizal-forming basidiomycete Amanita muscaria, trehalase activity was analysed from mycelium growing in liquid culture or on agar plates, and from mycorrhizas. Trehalase activity was found in both the culture medium and in mycelial extracts. The excreted trehalase was purified to apparent homogeneity by anion-exchange chromatography, gel filtration and preparative gel electrophoresis. The identified enzyme belongs to the group of acid trehalases. The apparent molecular mass of the native enzyme was estimated to be 165 kDa by gel filtration and 135 kDa by SDS–PAGE. Isoelectrofocusing indicated an isoelectric point (pI) of approx. 3.7, which is typical for acid trehalases. The enzyme is highly specific for its substrate trehalose, with an apparent Km of 0.38 mM. Validamycin and sucrose act as competitive inhibitors with Ki values of 45 nM and 15 mM, respectively. The activity of acid trehalase excreted into the growth medium is independent of the carbon source (glucose or trehalose), revealing that the enzyme is not regulated by its substrate trehalose. Nevertheless, fungal hyphae grown in the absence of an external carbon source showed enhanced enzyme excretion. The biochemical characteristics of the trehalase activity measured in the mycelium are in the same range as those determined for the excreted enzyme. The enzyme is localized in the cell wall and its activity is strongly decreased in ectomycorrhizas.