Incidence, size and spatial structure of clones in second-growth stands of coast redwood, Sequoia sempervirens (Cupressaceae)

The ecology and evolutionary potential of coast redwood (Sequoia sempervirens) is significantly influenced by the important role clonal spread plays in its reproduction and site persistence. In nine second-growth stands, amplified fragment length polymorphisms (AFLPs) were used to identify redwood clonal architecture. Clones (multistem genets) dominated sites by representing an average of 70% of stems measured, ranging in size from two to 20 stems. As a result, a relatively small number of genets can monopolize a disproportionate amount of site resources, are more likely to persist over time, and have greater on-site genetic representation. Clones were not limited to fairy-ring structures, but consisted of a wide range of shapes including concentric rings, ring chains, disjunct, and linear structures. Between-ramet distances of up to 40 m were measured, indicating that clonal reproduction is not limited to basal stump resprouting. Clonal structure in second-growth stands was similar to earlier reports from old growth, emphasizing the importance of site persistence and long-term, gradual site development. Smaller ramet numbers per genet in old growth is probably due to local within-genet self thinning. Management and conservation of redwoods will benefit from a better understanding of the dynamics and structure of clonal spread in these forests.     

Microsatellite markers in coast redwood (Sequoia sempervirens)

Five polymorphic microsatellite markers are described for coast redwood, a hexaploid conifer. Dinucleotide (three), trinucleotide (one) and tetranucleotide (one) repeat loci were isolated from a genomic plasmid library and amplified in a test population of 25 second‐growth redwoods. There were three to 11 alleles per locus and observed heterozygosities ranged from 0.00 to 0.80. The nonheterozygous locus (a CTTA repeat) is probably of chloroplast origin and may have potential as a marker for paternal inheritance.     

Shade-avoidance responses in two common coastal redwood forest species,Sequoia sempervirens (Taxodiaceae) and Satureja douglasii (Lamiaceae), occurring in various light quality environments

Shade-avoidance responses were examined for two species common to the coastal redwood forest, Sequoia sempervirens and Satureja douglasii. Sequoia seedlings demonstrated a shade-avoidance response when given end-of-day far-red light by increased hypocotyl, epicotyl, and first-node extension, and greater total number of needles and reduced anthocyanin concentration. Thus, Sequoia seedlings respond as sun-adapted plants. Satureja has several leaf monoterpene chemotypes that occur in different light environments including the redwood forest, and the types responded differently to the light treatments. The pulegone type responded to end-of-day far-red light as a sun-adapted plant with significant extension growth, increased leaf area and chlorophyll, and reduced anthocyanin. The isomenthone type responded as a shade-tolerant plant and did not exhibit extension growth nor a change in other parameters with end-of-day far-red light. However, the carvone and bicyclic types had variable responses depending on the parameter studied, which indicated genetic variation for these traits.     

The effects of nitrogen, light and water availability on tropic leaf movements in soybean (Glycine max)

Abstract. Rapid, tropic leaf movements and photo-synthetic responses of the heliotropic plant, soybean, Glycine max cv. Cumberland, grown under two different nitrogen, three different light and two different water treatments were examined. Measurements of leaf orientation during midday periods outdoors, and tropic reorientation of leaflets in response to vertical illumination indoors, revealed a positive, linear relationship between leaf water potential and the cosine of the angle of incidence between the leaf and the direct beam of the excitation light. This relationship was altered by nitrogen availability, such that a lower cosine of incidence (lower leaf irradiance) for a given leaf water potential was measured for plants grown under low nitrogen compared to those grown under high nitrogen. Additionally, plants grown under low nitrogen and low water availability showed more rapid rates of leaf movement compared to plants receiving high levels of these resources. Light regime during growth had no effect on the relationship between the cosine of incidence and leaf water potential. Reduced water and nitrogen availabilities during growth resulted in lower photosaturated rates of photosynthesis and stomatal conductance, as well as alterations in the relationship between these parameters. Thus, higher values for the ratio of intercellular CO₂/ambient CO₂ were measured for low-N grown plants (higher nitrogen use efficiencies) and lower values of this ratio for water stressed plants (higher water use efficiencies). The results show that environmental growth conditions other than water availability have the potential to modify leaf orientation responses to vectorial light in heliotropic legumes such as soybean. This has implications for the potential of heliotropic movements to minimize environmental stress-induced damage to the photosynthetic apparatus, and to modulate leaf-level resource use efficiencies.     

DNA variation in a conifer,Cryptomeria japonica (Cupressaceae sensu lato)

We investigated the nucleotide variation of a conifer, Cryptomeria japonica, and the divergence between this species and its closest relative, Taxodium distichum, at seven nuclear loci (Acl5, Chi1, Ferr, GapC, HemA, Lcyb, and Pat). Samples of C. japonica were collected from three areas, Kantou-Toukai, Hokuriku, and Iwate. No apparent geographic differentiation was found among these samples. However, the frequency spectrum of the nucleotide polymorphism revealed excesses of intermediate-frequency variants, which suggests that the population was not panmictic and a constant size in the past. The average nucleotide diversity, pi, for silent sites was 0.00383. However, values of pi for silent sites vary among loci. Comparisons of polymorphism to divergence among loci (the HKA test) showed that the polymorphism at the Acl5 locus was significantly lower. We also observed a nearly significant excess of replacement polymorphisms at the Lcyb locus. These results suggested possibilities of natural selection acting at some of the loci. Intragenic recombination was detected only once at the Chi1 locus and was not detected at the other loci. The low level of population recombination rate, 4Nr, seemed to be due to both low level of recombination, r, and small population size, N.     

The ethnobotany,leaf anatomy,essential oil variation and biological activity of Pteronia incana (Asteraceae)

The only available ethnobotanical information on Pteronia incana has been recorded by the Montagu Museum in the Western Cape Province of South Africa. It was reported that the plant is used to treat influenza, fever, kidney ailments and backache. In common with other species of Pteronia, the plant contains an essential oil reminiscent of pine turpentine oil with β-pinene, limonene, 1,8-cineole, myrcene, spathulenol, p-cymene and methyleugenol as main compounds present in all or most of the samples, with smaller amounts of α-pinene, sabinene, γ-terpinene, terpinen-4-ol, biclogermacrene, globulol and α-bisabolol in some of the distillates. We investigated the oil composition of 11 individual plants collected at three geographically distant localities but found limited variation, both within and between populations. Leaf sections of P. incana showed that it is anatomically similar to P. divaricata in the presence of a secretory duct associated with the main vascular bundle (and often other bundles as well), in addition to glandular and non-glandular trichomes on both leaf surfaces. One yeast (Cryptococcus neoformans), two Gram-negative bacteria (Moraxella catarrhalis and Klebsiella pneumoniae) and one Gram-positive bacterium (Mycobacterium smegmatis) were selected for antimicrobial activity studies using the minimum inhibitory concentration (MIC) microtitre plate method. The results showed that methanol:dichloromethane (MeOH:CH₂Cl₂) extracts were active against M. smegmatis (lowest MIC values of 0.5–0.8 mg/ml) and C. neoformans (lowest MIC values of 0.5–2.0 mg/ml). The essential oil was most active against C. neoformans (lowest MIC value of 0.3 mg/ml). These results provide a scientific rationale for the use of P. incana in Cape herbal medicine.     

Hydraulic efficiency and safety of branch xylem increases with height in Sequoia sempervirens (D. Don) crowns

The hydraulic limitation hypothesis of Ryan & Yoder (1997, Bioscience 47, 235-242) suggests that water supply to leaves becomes increasingly difficult with increasing tree height. Within the bounds of this hypothesis, we conjectured that the vertical hydrostatic gradient which gravity generates on the water column in tall trees would cause a progressive increase in xylem 'safety' (increased resistance to embolism and implosion) and a concomitant decrease in xylem 'efficiency' (decreased hydraulic conductivity). We based this idea on the historically recognized concept of a safety-efficiency trade-off in xylem function, and tested it by measuring xylem conductivity and vulnerability to embolism of Sequoia sempervirens branches collected at a range of heights. Measurements of resistance of branch xylem to embolism did indeed show an increase in 'safety' with height. However, the expected decrease in xylem 'efficiency' was not observed. Instead, sapwood-specific hydraulic conductivities (Ks) of branches increased slightly, while leaf-specific hydraulic conductivities increased dramatically, with height. The latter could be largely explained by strong vertical gradients in specific leaf area. The increase in Ks with height corresponded to a decrease in xylem wall fraction (a measure of wall thickness), an increase in percentage of earlywood and slight increases in conduit diameter. These changes are probably adaptive responses to the increased transport requirements of leaves growing in the upper canopy where evaporative demand is greater. The lack of a safety-efficiency tradeoff may be explained by opposing height trends in the pit aperture and conduit diameter of tracheids and the major and semi-independent roles these play in determining xylem safety and efficiency, respectively.     

The contribution of fog to the water relations of Sequoia sempervirens (D. Don): foliar uptake and prevention of dehydration

Fog is a defining feature of the coastal California redwood forest and fog inputs via canopy drip in summer can constitute 30% or more of the total water input each year. A great deal of occult precipitation (fog and light rain) is retained in redwood canopies, which have some of the largest leaf area indices known (Westman & Whittaker, Journal of Ecology 63, 493–520, 1975). An investigation was carried out to determine whether some fraction of intercepted fog water might be directly absorbed through leaf surfaces and if so, the importance of this to the water relations physiology of coast redwood, Sequoia sempervirens. An array of complimentary techniques were adopted to demonstrate that fog is absorbed directly by S. sempervirens foliage. Xylem sap transport reversed direction during heavy fog, with instantaneous flow rates in the direction of the soil peaking at approximately 5–7% of maximum transpiration rate. Isotopic analyses showed that up to 6% of a leaf's water content could be traced to a previous night's fog deposition, but this amount varied considerably depending on the age and water status of the leaves. Old leaves, which appear most able to absorb fog water were able to absorb distilled water when fully submersed at an average rate of 0.90 mmol m² s^?1, or about 80% of transpiration rates measured at the leaf level in the field. Sequoia sempervirens has poor stomatal control in response to a drying atmosphere, with rates of water loss on very dry nights up to 40% of midday summer values and rates above 10% being extremely common. Owing to this profligate water use behaviour of S. sempervirens, it appears that fog has a greater role in suppressing water loss from leaves, and thereby ameliorating daily water stress, than in providing supplemental water to foliar tissues per se. Although direct foliar absorption from fog inputs represents only a small fraction of the water used each day, fog's in reducing transpiration and rehydrating leaf tissues during the most active growth periods in summer may allow for greater seasonal carbon fixation and thus contribute to the very fast growth rates and great size of this species.     

A comparison of leaf physiology and anatomy of Quercus (section Erythrobalanus-Fagaceae) species in different light environments

Physiological and anatomical attributes of leaves were examined of three species of Quercus section Erythrobalanus. All three species occur in moist temperate deciduous forests of eastern North America. Seedlings of each species were grown in different light conditions for comparison. The attributes measured were net photosynthesis, stomatal conductivity, blade and cuticle thickness, stomatal density, thickness of upper and lower epidermis, and thickness of palisade mesophyll. The results generally demonstrate the close association between anatomical adaptations and efficiency of physiological processes; they also elucidate the distribution patterns of the three Quercus species across the forest topography in southern New England. The most drought-tolerant and light-demanding species, Q. velutina (Lam.), exhibited the greatest leaf anatomical plasticity, the highest net photosynthesis in the different light conditions, and the lowest stomatal area per unit area of leaf. The most drought-intolerant species, Q. rubra (L.), exhibited the least leaf anatomical plasticity, the lowest net photosynthesis in the different light conditions, and the highest stomatal area per unit area of leaf. Quercus coccinea (Muenchh.) usually exhibited values that were intermediate between Q. rubra and Q. velutina.     

Character displacement drives floral variation in Pelargonium (Geraniaceae) communities

Interactions between plant community members are an underexplored driver of angiosperm floral variation. We investigate character displacement as a potential contributor to floral variation in Pelargonium communities. Pelargoniums all place pollen on the ventral sides of their pollinators, potentially leading to interspecific pollen transfer (IPT) in sympatry. We show that the positions of pollen placement and receipt are determined by anther and style exsertion lengths. Using field experiments, we demonstrate that heterospecific species experience higher IPT if they have similar style lengths than when they have greater style length differences. Using crosses, we show that IPT has negative consequences on seed set. In combination, these results suggest that character displacement in style length is likely to reduce IPT and increase female fitness in sympatry. Patterns of style length variation across 29 different Pelargonium communities suggest that character displacement has occurred in multiple communities. Furthermore, analyses using a wide-ranging species pair show that style lengths are more different between sympatric populations than they are between allopatric populations. In addition to pollinators as agents of floral divergence, this study suggests that variation in Pelargonium community structure has driven style length variation through character displacement.     

Chimeric patterns in Juniperus chinensis 'Torulosa Variegata' (Cupressaceae) expressed during leaf and stem formation

Juvenile leaves of the variegated Hollywood juniper, Juniperus chinensis 'Torulosa Variegata', have sectorial chimeras of variable widths and lengths. Sectors extend over several nodes often as small as 1/24 the circumference of the leaf. Other chimeras appear as light green to yellow streaks but are actually internal, dark green corpus sectors often occupying less than 1/20 of the cross sectional area of a leaf. On the basis of the sizes of these two types of sectors, there seems to be ideally about 168 founder cells comprising 63 tunica cells and 105 corpus cells; 49 of the latter are contiguous with the tunica and 66 are located deeper in the corpus. Similarly, sectoring in axillary branches of original chimeric sprays have the same types of sectoring. It is hypothesized that the outer rings of founder cells form two arcs of 12 cells around the stem apex, one for each of two leaves at a node of the decussate shoot, of a circumference of about 50 cells.     

Comparative leaf anatomy of Alpinia species (Zingiberaceae) in Malaysia

The leaf anatomy of 22 Alpinia species from the Malay Peninsular was investigated. Results show that a combination of characters such as outlines of midribs, petioles, leaf margins and their relative sizes in transverse sections as well as the presence of hypodermal fibres in the lamina, midribs and petioles can be used in the identification of species. All four species of subsection Allughas possess similar characters such as the presence of abaxial and adaxial fibres in the midrib and petioles and the absence of hypodermis in the lamina while the reverse is true for all species of subsection Catimbium. Alpinia conchigera and A. galanga appear to be closely related while A. oxymitra is distinct from all the species studied.     

Partitioning of leaf nitrogen with respect to within canopy light exposure and nitrogen availability in peach (Prunus persica)

Summary Relationships between leaf nitrogen content and within canopy light exposure were studied in mature nectarine peach trees (Prunus persica cv. Fantasia) that had received 0, 112, 196, 280 or 364 kg of fertilizer nitrogen per hectare per year for the previous 3 years. The relationships between light saturated leaf CO₂ assimilation rates and leaf nitrogen concentration were also determined on trees in the highest and lowest nitrogen fertilization treatments. The slope of the linear relationship between leaf N content per unit leaf area and light exposure was similar for all nitrogen treatments but the y-intercept of the relationship increased with increasing N status. The slope of the relationship between leaf N content per unit leaf area and light saturated CO₂ assimilation rates was greater for the high N trees than the low N trees, but maximum measured leaf CO₂ assimilation rates were similar for both the high and low N treatments. A diagrammatic model of the partitioning of leaf photosynthetic capacity with respect to leaf light exposure for high and low nitrogen trees suggests that the major influence of increased N availability is an increase in the photosynthetic capacity of partially shaded leaves but not of the maximum capacity of highly exposed leaves.     

The comparative leaf anatomy of Agave, Beschorneria, Doryanthes and Furcraea species (Agavaceae: Agaveae)

The leaf anatomy is compared of 35 ppecies and one variety of Agave , three species of Furcraea , one species of Beschorneria and one species of Doryanrhes , four genera assigned to the tribe Agaveae by Hutchinson in his classification of the family Agavaceae. The genera can be readily differentiated by leaf anatomy. There are close similarities between Agave, Beschorneria and Furcraea , but Doryanthes differs widely from these genera. Sufficient anatomical differences exist to differentiate the Agave species examined.     

C3 and C4 leaf anatomy types in Camphorosmeae (Camphorosmoideae,Chenopodiaceae)

Complementary to our previous project on the molecular phylogeny of Camphorosmeae, the leaf anatomy of ca. 35 species including all non-Australian and selected Australian species was studied by use of light microscopy. Nine anatomical leaf types were described, compared to previous classifications, and discussed with regard to their putative evolution on the background of phylogenetic trees. Particular emphasis was given to the relationships between the C₃ and C₄ leaf types: Chenolea type (C₃), Eokochia type (C₃), Neokochia type (C₃), Sedobassia type (C₃/C₄ intermediate), Bassia prostrata type (C₄), B. muricata type (C₄), B. eriantha type, B. lasiantha type (C₄), Camphorosma type (C₄). The main results and conclusions were: (1) Two unusual new C₃ leaf types: Chenolea with microfenestrate chlorenchyma, Eokochia with unique complex vascular bundles; (2) Sedobassia interpreted as anatomically C₃/C₄ intermediate by kranz-like bundle sheath cells is the first C₃/C₄ intermediate in Camphorosmeae and found in a derived position; (3) Neokochia type detected as the likely starting point for all four C₄ leaf types and for the C₃/C₄ intermediate; (4) hypodermis of C₄ types originated from outermost chlorenchyma layer of C₃ types and lost multiple times during further evolution; (5) atriplicoid Bassia. lasiantha type without water storage tissue evolved from kochioid B. muricata type; (6) two independent gains of C₄ photosynthesis, one in Bassia and one in Camphorosma; (7) depending on the lineage, leaf architecture remains comparatively stable (Australian Camphorosmeae) or shows an unexpected plasticity (Bassia scoparia group).     

Saturating light and not increased carbon dioxide under ocean acidification drives photosynthesis and growth in Ulva rigida (Chlorophyta)

Carbon physiology of a genetically identified Ulva rigida was investigated under different CO_2(aq) and light levels. The study was designed to answer whether (1) light or exogenous inorganic carbon (Ci) pool is driving growth; and (2) elevated CO_2(aq) concentration under ocean acidification (OA) will downregulate CA_ext-mediated dehydration and alter the stable carbon isotope (δ¹³C) signatures toward more CO₂ use to support higher growth rate. At pH_T 9.0 where CO_2(aq) is <1 μmol L⁻¹, inhibition of the known use mechanisms, that is, direct uptake through the AE port and CA_ext-mediated dehydration decreased net photosynthesis (NPS) by only 56–83%, leaving the carbon uptake mechanism for the remaining 17–44% of the NPS unaccounted. An in silico search for carbon-concentrating mechanism elements in expressed sequence tag libraries of Ulva found putative light-dependent transporters to which the remaining NPS can be attributed. The shift in δ¹³C signatures from –22‰ toward –10‰ under saturating light but not under elevated CO_2(aq) suggest preference and substantial use to support photosynthesis and growth. U. rigida is Ci saturated, and growth was primarily controlled by light. Therefore, increased levels of CO_2(aq) predicted for the future will not, in isolation, stimulate Ulva blooms.     

Survey of leaf anatomy,especially secretory structures,of tribeCaesalpinieae (Leguminosae,Caesalpinioideae)

We studied leaflet anatomy, emphasizing secretory structures, from herbarium specimens of 128 species of 44 genera of tribeCaesalpinieae, using clearings, resin sections, and scanning electron microscopy. These observations, combined with those from our three earlier papers, provide a survey of 210 species representing all genera. Seventy-three species had secretory structures: 21 had glands or gland-like trichomes, 40 had living mesophyll idioblasts, and nine had cavities (three species each had two different types). Five additional species, all inCercidium (Caesalpinia group), had paired or clustered large spheroidal, thick-walled, empty cells (veinlet idioblasts) interconnected by perforation plate-like gaps. Secretory structures have systematic significance at various taxonomic levels.     

Systematic vegetative anatomy and ensiform leaf development in Xyris (Xyridaceae)

Ensiform leaf development in monocotyledons follows a broadly similar sequence in a wide range of relatively unrelated taxa, indicating a plastic developmental pattern, possibly associated with stressed environmental conditions, since Xyris species tend to grow in relatively damp but nutrient-poor environments. The bifacial leaf sheath surrounds the apex and the subadjacent primordium. A conical unifacial leaf tip 'Vorläuferspitze' is established at an early stage, followed by extension growth in the region behind it, generating a unifacial ensiform blade. Root and rhizome structure are also described in a systematic context, particularly in comparison with related taxa in Xyridaceae and other commelinoid monocotyledons, although information on these structures is relatively sparse.     

The role of leaf and canopy-level gas exchange in the replacement of Quercus virginiana (Fagaceae) by Juniperus ashei (Cupressaceae) in semiarid savannas

Photosynthesis, transpiration, and leaf area distribution were sampled in mature Quercus virginiana and Juniperus ashei trees to determine the impact of leaf position on canopy-level gas exchange, and how gas exchange patterns may affect the successful invasion of Quercus communities by J. ashei. Sampling was conducted monthly over a 2-yr period in 12 canopy locations (three canopy layers and four cardinal directions). Photosynthetic and transpiration rates of both species were greatest in the upper canopy and decreased with canopy depth. Leaf photosynthetic and transpiration rates were significantly higher for Q. virginiana (4.1–6.7 μmol CO₂·m⁻²·s⁻¹ and 1.1–2.1 mmol H₂O·m⁻²·s⁻¹) than for J. ashei (2.1–2.8 μmol CO₂·m⁻²·s⁻¹ and 0.7–1.0 mmol H₂O·m⁻²·s⁻¹) in every canopy level and direction. Leaves on the south and east sides of both species had higher gas exchange rates than leaves on the north and west sides. Although Quercus had a greater mean canopy diameter than Juniperus (31.3 vs. 27.7 m²), J. ashei had significantly greater leaf area (142 vs. 58 m²/tree). A simple model combining leaf area and gas exchange rates for different leaf positions demonstrated a significantly greater total canopy carbon dioxide uptake for J. ashei compared to Q. virginiana (831 vs. 612 g CO₂·tree⁻¹·d⁻¹, respectively). Total daily water loss was also greater for Juniperus (125 vs. 73 Ltree⁻¹·d⁻¹). Differences in leaf gas exchange rates were poor predictors of the relationship between the invasive J. ashei and the codominant Q. virginiana. Leaf area and leaf area distribution coupled with leaf gas exchange rates were necessary to demonstrate the higher overall competitive potential of J. ashei.