BRANCHING PATTERNS OF COLUMNAR CACTI: INFLUENCES ON PAR INTERCEPTION AND CO2 UPTAKE

Field measurements and a computer model were used to determine how stem shape and arrangement of stems in space affect interception of photosynthetically active radiation (PAR) and CO₂ uptake under otherwise optimal conditions for four species of columnar cacti (Carnegiea gigantea, Lophocereus schottii, Pachycereus pringlei, and Stenocereus thurberi). In simulations where the number of widely spaced stems was increased from 1 to 19 but plant volume remained constant, surface area and PAR interception increased, leading to 3-fold increases in whole-plant CO₂ uptake. Increasing the distance between stems from 0 cm to infinity decreased self-shading and increased predicted CO₂ uptake 4-fold. Stem length, diam, ribbing characteristics, and spine coverage also influenced PAR interception. The model indicated that the observed higher frequency of branches on the south side of the trunk of C. gigantea had only a slight, though positive, effect on CO₂ uptake for single-branched plants. Because of its greater surface area (A), a five-stemmed plant of C. gigantea typical for a field site near Tucson, Arizona was predicted to have 52% more CO₂ uptake than a single-stemmed plant of the same volume (V). Although large A/V decreases water storage per unit transpiring area, whole-plant CO₂ uptake can be increased when A/V is increased by branching for these constant-volume plants. However, the stems must be arranged to avoid excessive self-shading and thus keep the area below PAR compensation small.     

LEAF DEVELOPMENT AND CROWN GEOMETRY OF TWO IRIARTEOID PALMS

We examined changes in pinnately compound leaf morphology and crown geometry that occur during height growth of the iriarteoid palms Socratea exorrhiza and Iriartea deltoidea in tropical wet forest of Costa Rica. Although light availability increased with height, the number of leaves per plant was relatively constant. Total leaf area, however, was much larger in taller individuals. Increases in linear dimensions of leaves (length and width) was responsible for less than half of this greater surface area. More important was the transition from a basically dorsiventral display of leaflets in small individuals to a more radial display in taller plants. Production of leaflets in more than one plane resulted in leaves whose surface area was more than twice the horizontally projected area and whose lateral light interception was greatly enhanced. S. exorrhiza, a faster-growing and more light-demanding species, undergoes this transformation in leaf morphology at heights between 3 and 6 m; whereas in the slower-growing and more shade-tolerant I. deltoidea this occurs at heights between 10 and 20 m. Pinnate leaves, with dense radial packing of leaflets along the rachis, are functionally comparable to branches of dicotyledonous trees and may have been important for the evolution of arborescence in palms.     

A comparison of pectoral fin ray morphology and its impact on fin ray flexural stiffness in labriform swimmers

The organization of tissues in appendages often affects their mechanical properties and function. In the fish family Labridae, swimming behavior is associated with pectoral fin flexural stiffness and morphology, where fins range on a continuum from stiff to relatively flexible fins. Across this diversity, pectoral fin flexural stiffness decreases exponentially along the length of any given fin ray, and ray stiffness decreases along the chord of the fin from the leading to trailing edge. In this study, we examine the morphological properties of fin rays, including the effective modulus in bending (E), second moment of area (I), segmentation, and branching patterns, and their impact on fin ray stiffness. We quantify intrinsic pectoral fin ray stiffness in similarly sized fins of two closely related species that employ fins of divergent mechanics, the flapping Gomphosus varius and the rowing Halichoeres bivittatus. While segmentation patterns and E were similar between species, measurements of I and the number of fin ray branch nodes were greater in G. varius than in H. bivittatus. A multiple regression model found that of these variables, I was always significantly correlated with fin ray flexural stiffness and that variation in I always explained the majority of the variation in flexural stiffness. Thus, while most of the morphological variables quantified in this study correlate with fin ray flexural stiffness, second moment of area is the greatest factor contributing to variation in flexural stiffness. Further, interspecific variation in fin ray branching pattern could be used as a means of tuning the effective stiffness of the fin webbing to differences in swimming behavior and hydrodynamics. The comparison of these results to other systems begins to unveil fundamental morphological features of biological beams and yields insight into the role of mechanical properties in fin deformation for aquatic locomotion.     

Patterns of Primary Succession on the Foreland of Coleman Glacier,Washington, USA

Patterns of community development vary among studied glacier forelands around the world. However, there have been few studies of primary succession on glacial forelands in temperate regions of North America. We described patterns in community composition, vegetation cover, diversity, and vegetation heterogeneity during primary succession on the foreland of Coleman Glacier, in Washington State, USA. Community composition changed rapidly with high turnover between age classes. Cover increased through succession as expected. Species richness and diversity were highest in early succession at small scales and in late succession at larger scales. At small scales, heterogeneity decreased in early succession but increased in mature sites. At larger scales, heterogeneity reached its lowest point earlier in succession. These scale-dependent patterns in diversity and heterogeneity differ from results of other studies of glacier forelands. We hypothesize that these patterns arise due to the development of a dense canopy of the deciduous shrub Alnus viridis followed by a dense canopy of Abies amabilis, Tsuga heterophylla, and Pseudotsuga menziesii.     

THE EFFECT OF SEED DIMORPHISM ON THE GERMINATION AND SURVIVAL OF SALICORNIA EUROPAEA L. POPULATIONS

Seed dimorphism influenced the germination behavior of Salicornia europaea L., with small seeds being more dormant and less salt tolerant than large seeds. All of the large less dormant seeds of S. europaea germinated prior to May, and all seedlings produced after this time were from small seeds. A persistent seed bank was maintained by the small dormant seeds. Survivorship was relatively constant during the normal germination season, but increasing salinity stress at any time during the spring reduces the chance of seedling survival. Little germination occurred from July through September because of the high surface soil salinities during this period.     

Competition between Salicornia europaea and Atriplex prostrata (Chenopodiaceae) along an experimental salinity gradient

Salicornia europaea L. is a halophyte that often occupies the lowestand most saline (>3.5% total salt) areas of salt marshes. Atriplexprostrata Boucher is less salt tolerant than S. europaea and oftengrows in a less saline (<2.0% total salts) zone adjacent to S. europaea. The purpose of this experiment was to determine thecompetitive outcome when these two species are grown at differentsalinities to ascertain the extent salinity and competition affect plantzonation. Plants were grown in a de Wit replacement series at 85, 170,and 340 mM NaCl in half-strength Hoagland's no. 2 nutrient solution fortwo months. There was a significant effect of salt concentration,competition, and their interaction on biomass production of S. europaea plants. However, only salt concentration significantly affectedbiomass production of A. prostrata plants. Results of thisexperiment confirmed the results of other studies that demonstrated thatthe more salt tolerant species were less competitive at lower salinities. Atriplex prostrata was the better competitor at 85 mM NaCl, whereasS. europaea was the better competitor at 340 mM NaClbecause growth of A. prostrata was inhibited. At 170 mMNaCl, A. prostrata biomass production decreased more than S. europaea biomass in mixed culture.     

BRANCH GEOMETRY AND EFFECTIVE LEAF AREA: A STUDY OF TERMINAUA-BRANCHING PATTERN. 1. THEORETICAL TREES

Single lateral branches and branch tiers of Terminalia catappa L. are simulated and drawn by computer. Leaf clusters on the branches are approximated by discs, and the effective leaf areas are determined by use of Dirichlet domains. Theoretical optimal branching angles which produce the maximum effective leaf area are obtained from simulations. Symmetrical and asymmetrical branching angles are contrasted; the latter characterize real trees. Varying leaf disc radius and ratio of branch-unit lengths affects optimal branching angles, as does the symmetry of a tier of five branches. Leaf area indices for individual branches and branch tiers are given for all simulations. The number of branches in a tier has a major effect on leaf area index and effective leaf area. The theoretical optimal branching angles of many simulations are very close to the values observed in real trees of T. catappa. We conclude that the observed branching angles and number of branches in a tier of this species optimize light interception within constraints of a fixed pattern of branching, one that is widespread among tropical trees.     

Relationships between growth,population structure and sea surface temperature in the temperate solitary coral <Emphasis Type="Italic">Balanophyllia europaea</Emphasis> (Scleractinia,Dendrophylliidae)

The demographic characteristics of the solitary zooxanthellate scleractinian Balanophyllia europaea, endemic to the Mediterranean, were determined in six populations, on a latitudinal gradient along the Italian coast, and compared with the mean annual sea surface temperature (SST). Growth rate correlated negatively, and asymptotic length of the individuals positively with SST. With increasing SST, the distributions of age frequencies moved away from a typical steady state structure (i.e., exponential decrease in the frequency of individuals with age), indicating less stable populations and showed a deficiency of individuals in the younger-age classes. These observations suggest that high temperatures are an adverse factor to the B. europaea symbiosis. Using projected increases in seawater temperature, most of the B. europaea populations in the Mediterranean are expected to be close to their thermal limits by 2100 and the populations at that time may support few young individuals. Communicated by Environment Editor Prof. van Woesik.     

Pattern of morphological variation of Salicornia in north Europe

Among the few available morphological traits in the genus Salicornia L. (Amaranthaceae/Chenopodiaceae), most are extremely variable within species probably due to high levels of plasticity. In addition, identifying Salicornia taxa is further complicated by that these plants lose many of their diagnostic characters upon drying. Morphological studies of fresh (or liquid preserved) specimens is thus important in taxonomical studies. The objective of this study was a numerical morphological analyses of Salicornia populations in the Nordic countries and an adjacent Russian region with the aim to ascertain whether taxonomic division of this genus based on morphology is feasible. In all, 666 plants were collected for morphometric measurements from 31 localities and 52 populations or subpopulations of the North Sea coasts, Danish straits, Baltic Sea, the Norwegian Sea, Barents Sea, and the White Sea areas. For practical reasons, part of the samples was studied fresh, and part preserved in FAA pending measurements. Data were analyzed by principal component analysis (PCA) and hierarchical cluster analysis. The main taxonomical division was found between the diploid S. europaea s.l. and the tetraploid S. procumbens Sm., and was mainly based on fertile segment and flower morphology characteristics. Data also show ecological differences: diploids inhabit the upper part of salt marshes, whereas tetraploids grow in the hydrolittoral zone and are restricted to areas of regular tidal influence. It was not possible to make a morphological distinction between the two cryptic diploid species S. europaea s.s. and S. perennans Willd. in the present study. North Norwegian S. europaea clearly deviated from more southern populations, but taxonomical conclusions based only on morphology were avoided. In contrast, the division of S. procumbens into two geographical races/subspecies, the southern subsp. procumbens and the northern subsp. pojarkovae was supported.     

Salt stress effects on growth,pigments, proteins and lipid peroxidation in <Emphasis Type="Italic">Salicornia persica</Emphasis> and <Emphasis Type="Italic">S. europaea</Emphasis>

The effects of NaCl stress on growth, water status, contents of protein, proline, malondialdehyde (MDA), various sugars and photosynthetic pigments were investigated in seedlings of Salicornia persica and S. europaea grown in vitro. Seeds were germinated under NaCl (0, 100, 200, 300, 400, 500 and 600 mM) on Murashige and Skoog medium for 45 d. The shoot growth of both species increased under low NaCl concentration (100 mM) and then decreased with increasing NaCl concentrations. In contrast to S. persica, root length in S. europaea reduced steadily with an increase in salinity. Proline content in S. persica was higher than in S. europaea at most NaCl concentrations. Proline, reducing saccharide, oligosaccharide and soluble saccharide contents increased under salinity in both species. In contrast, contents of proteins and polysaccharides reduced in both species under salt stress. MDA content remained close to control at moderate NaCl concentrations (100 and 200 mM) and increased at higher salinities. MDA content in S. europaea was significantly higher than S. persica at higher salinities. Salt treatments decreased K⁺ and P contents in seedlings of both species. Significant reduction in contents of chlorophylls and carotenoids due to NaCl stress was also observed in seedlings of both species. Some differences appeared between S. persica and S. europaea concerning proteins profile. On the basis of the data obtained, S. persica is more salt-tolerant than S. europaea.     

MECHANICAL PROPERTIES WITHIN THE GROWTH ZONE OF CORN ROOTS INVESTIGATED BY BENDING EXPERIMENTS I. PRELIMINARY OBSERVATIONS

Bending experiments were performed to analyze mechanical properties within the apical 15 mm of the primary root of corn, Zea mays. Force required to maintain a root in a bent shape declined with the logarithm of time, indicating significant stress relaxation. Spatial distributions of local curvature, strain due to bending, and geometrical moment of inertia were calculated from digitized photographs of mechanically bent roots during and after imposition of a bending moment. When roots were bent to nonuniform curvature (maximum K = 3 cm^–1 at the location 8 mm from the root tip), “irreversible” curvature (residual curvature 4 hr after release of the bending moment) was approximately 44% of initial curvature. This percentage of irreversible curvature (and associated bending strain) was uniform throughout the growth zone. When roots were bent to uniform curvature of either 1 cm^–1 or 1.67 cm^–1, “springback” curvature (residual curvature after two minutes of bending) varied from 60% of initial curvature in the apical regions (at 4 mm) to 85% of initial curvature at the base of the growth zone. The results are consistent with a model in which total strain is proportional to stress, and irreversible strain at a particular location is proportional to total strain at the same location.     

ARCHITECTURE AND CROWN GEOMETRY IN TABEBUIA ROSEA (BIGNONIACEAE)

In the tropical tree Tabebuia rosea, seedlings form an erect unbranched stem with rhythmic growth. Three distinct and predictable geometrical stages are then recognized in crown development. Sympodial branching begins with the arrest of the terminal bud of the trunk and symmetric outgrowth of a pair of subtending lateral buds. During an intermediate phase, branching becomes asymmetric at about Order 5. At each sympodial bifurcation there is differentiation between vigorous, relatively straight main branches (leaders) and less vigorous laterals forming regular pseudomonopodial branch complexes, which collectively constitute the cup-shaped crown. Finally, dormant lateral buds in the lowest bifurcation of the trunk are released and reiterate the original crown form. Ultimately an erect, apparently monopodial tree is formed by a set of superimposed cup-shaped crowns. Crown development of Tabebuia is unique because it involves predictable ontogenic changes in branching patterns. Crowns of open-grown Tabebuia consist of relatively few, wide branch tiers, crowns of forest grown trees are tall and narrow. Analysis of the adaptive geometry of wide vs. narrow crowns through computer simulation illustrates the precise cost of mechanical support for terminal leaf rosettes at successive developmental phases and suggests that tall, erect, narrow, and multi-tiered crowns are more efficient than wide open crowns.     

Three dimensional typological studies using scanning electron microscopy for characterization of Termitomyces pellets obtained from submerged growth conditions

There are gaps in existing understanding of fungal pellet growth dynamics. We used scanning electron microscopy (SEM) for morphological characterization of the biomass organization of Termitomyces pellets for seven species: T. microcarpus (TMI1), T. albuminosus (TAL1, TAL2), T. striatus (TSTR), T. aurantiacus (TAUR), T. heimii (THE1, THE2), T. globulus (TGLO) and T. clypeatus (TCL1, TCL2, TCL3, TCL4, TCL5). We assessed the utility of SEM for morphological and structural characterization of Termitomyces spp. in three dimensional (3D) pellet form to identify ideal pellet morphology for industrial use. Typological classification of Termitomyces species was based on furrows, isotropy, total motifs and fractal dimensions. The pellets formed were entangled and exhibited highly compacted mycelial mass with microheterogeneity and microporosity. The mean density of furrows of Termitomyces species was between 10,000 and 11,300 cm/cm², percentage isotropy was 30?80 and total motifs varied from 300 to 2500. TGLO exhibited the highest furrow mean density, 11243 cm/cm², which indicated a compact, cerebroid structure with complex ridges and furrows, whereas TAL2 exhibited the lowest furrow density. TMI1a exhibited a high percentage isotropic value, 74.6, TSTR exhibited the lowest, 30.9. Total motif number also was used as a typological classification parameter. Fractal values were 2.64?2.78 for various submerged conditions of Termitomyces species. TAL1 exhibited the highest fractal dimension and TAL2 the lowest, which indicates the complexity of branching patterns. Three-dimensional SEM image analysis can provide insight into pellet micromorphology and is a powerful tool for exploring topographical details of pellets.     

The expression of light-related leaf functional traits depends on the location of individual leaves within the crown of isolated Olea europaea trees

Background The spatial arrangement and expression of foliar syndromes within tree crowns can reflect the coupling between crown form and function in a given environment. Isolated trees subjected to high irradiance and concomitant stress may adjust leaf phenotypes to cope with environmental gradients that are heterogeneous in space and time within the tree crown. The distinct expression of leaf phenotypes among crown positions could lead to complementary patterns in light interception at the crown scale.Methods We quantified eight light-related leaf traits across 12 crown positions of ten isolated Olea europaea trees in the field. Specifically, we investigated whether the phenotypic expression of foliar traits differed among crown sectors and layers and five periods of the day from sunrise to sunset. We investigated the consequences in terms of the exposed area of the leaves at the tree scale during a single day.Key Results All traits differed among crown positions except the length-to-width ratio of the leaves. We found a strong complementarity in the patterns of the potential exposed area of the leaves among day periods as a result of a non-random distribution of leaf angles across the crown. Leaf exposure at the outer layer was below 60 % of the displayed surface, reaching maximum interception during morning periods. Daily interception increased towards the inner layer, achieving consecutive maximization from east to west positions within the crown, matching the sun’s trajectory.Conclusions The expression of leaf traits within isolated trees of O. europaea varies continuously through the crown in a gradient of leaf morphotypes and leaf angles depending on the exposure and location of individual leaves. The distribution of light-related traits within the crown and the complementarity in the potential exposure patterns of the leaves during the day challenges the assumption of low trait variability within individuals.     

Aggregative responses of brent geese on salt marsh and their impact on plant community dynamics

The aggregative responses and habitat preferences of a generalist herbivore, the dark-bellied brent goose Branta bernicla bernicla, feeding on salt marshes are examined in relation to vegetation community characteristics and the abundances of individual plant species. In the autumn, feeding was strongly concentrated on the low marsh, which had the highest biomass of the preferred food plant, Salicornia europaea. There was a strong aggregative response of the geese to the abundance of S. europaea. A decline in the availability of S. europaea led to an increase in the pattern of aggregation in relation to the two other major food plants on the low marsh, Aster tripolium and Puccinellia maritima. The availability of these food plants, however, reached critically low levels in mid-winter and the geese abandoned the low marsh for the high marsh. Within the high marsh, the plant communities selected tended to be dominated by the inedible species Limonium vulgare. The food plants selected were P. maritima in the winter and P. maritima and Triglochinmaritimum in the spring. On the high marsh, aggregative responses were shown to both P. maritima and T. maritimum, but in both cases, aggregation increased up to a critical level of biomass, and then declined. The prevention of grazing with exclosures for 3 years led to an increase in the abundance of P. maritima on both high and low marshes. This change was slight on high marsh but pronounced on low marsh, where S. europaea showed a decrease in abundance in the exclosures over this time. The implications of the aggregative responses for the population dynamics of P. maritima and S. europaea are discussed. Received: 11 September 1997 / Accepted: 12 December 1997     

Sulphide tolerance in coastal halophytes

The effect of sulphide on the growth of several species of salt-marsh plants was investigated. Relative growth rates were significantly reduced in two upper-marsh species, Festuca rubra and Atriplex patula, and in the lower-marsh species Puccinellia maritima. However the growth of Salicornia europaea, a species frequently associated with sulphide-containing sediments, was unaffected. In a separate experiment the wide ranging halophyte Aster tripolium, also appeared to be tolerant of sulphide at a concentration frequently encountered in salt marshes. Sulphide pretreatment inhibited the activity of two metallo-enzymes, polyphenol oxidase and external phosphatase, in plants from the upper marsh, but had no effect on enzymes from P. maritima or S. europaea. The rate of respiration by root tissue was significantly reduced in all of the species investigated but whereas the uptake of ⁸⁶rubidium was markedly inhibited in the other three species, uptake by S. europaea showed a significant stimulation. Similarly, whereas sulphide-grown plants of F. rubra, A. patula and P. maritima had a considerably reduced tissue iron content, the total iron concentration in S. europaea tissues was comparable to that of the controls. When the sulphide-tolerant species A. tripolium was grown in sulphide-containing media there was no significant effect on the tissue concentration of any of the elements investigated. These results are discussed in relation to possible mechanisms of sulphide toxicity and resistance.     

腾格里沙漠人工固沙植被演替生物土壤结皮盖度对沙埋的响应

生物土壤结皮(简称结皮)的形成与发展是沙区固沙植被建设成功的重要标志之一,其盖度随固沙植被演替的变化趋势是表征该植被系统地表稳定性的一个直观性生态学指标。利用空间代时间的方法,以腾格里沙漠不同始植年代(1956、1973、1981和1987年)固沙植被区发育的4种优势结皮-真藓(Bryum argenteum Hedw.)结皮、土生对齿藓(Didymodon vinealis(Brid.)Zand.)结皮、齿肋赤藓(Syntrichia caninervis Mitt.)结皮和藻-地衣-藓类混生结皮(Mixed crust)为研究对象,在测定结皮盖度、高度以及粗糙度随沙埋厚度逐渐增加变化的基础上,计算了使结皮盖度(从99.99%)开始降低的初始沙埋厚度(D1)和盖度降低为0%的临界沙埋厚度(D2),研究了该区固沙植被演替过程中结皮盖度对沙埋厚度增加的响应特征及其相关机制。结果显示:(1)4种结皮的盖度随沙埋厚度的增加呈logistic曲线逐渐降低。(2)在同一年代固沙区不同种间比较,混生结皮的D1值最小,D2值最大;真藓结皮的D1和D2值均小于其他两种藓类结皮;土生对齿藓结皮和齿肋赤藓结皮因固沙年限的不同而不同,在1956年固沙区,土生对齿藓结皮齿肋赤藓结皮,而在1973年固沙区和1981年固沙区,齿肋赤藓结皮土生对齿藓结皮。(3)随固沙植被演替,4种结皮盖度随沙埋厚度增加的降低速率逐渐减小,D1和D2值逐渐增大。(4)结皮总盖度随沙埋厚度增加的降低速率逐渐减小,表明随固沙植被演替,结皮对沙埋的抵御能力逐渐增强,固沙区植被系统地表稳定性增加。(5)结皮的粗糙度和高度随固沙植被演替逐渐增加,并显著影响了4种结皮盖度随沙埋厚度增加的降低速率、D1及D2值。研究为全面评估沙埋对沙区结皮结构、功能的影响乃至固沙植被稳定性提供了参考。     

LEAF DISPLAY,CANOPY STRUCTURE,AND LIGHT INTERCEPTION OF TWO UNDERSTORY PALM SPECIES

The implications of leaf size, leaf display, and crown size for whole-plant light interception were investigated in Geonoma cuneata and Asterogyne martiana, two understory palm species native to Central American rain forests. Adults of A. martiana had longer leaves, more leaves per plant, and greater total leaf area than G. cuneata adults. Geometric measurements within whole crowns were used to calculate light interception efficiency, a leaf-based measure of the proportion of total incident light that is intercepted by a crown. Light interception efficiency was higher in adult G. cuneata than in adult A. martiana; seedlings of the two species did not differ significantly in light interception efficiency. Decreased efficiency of adult A. martiana crowns was largely due to an increased proportion of pendent leaves. Compared to G. cuneata, adults of A. martiana had greater light interception capacity (the product of light interception efficiency and total leaf area), but they also had higher biomass costs of light interception. Lower biomass costs of light interception in adult G. cuneata enable this species to exploit successfully the most deeply shaded microsites in the rain forest understory.     

Growth and amino acid contents of<Emphasis Type="Italic">Spirulina platensis</Emphasis> with different nitrogen sources

The growth and amino acid contents of the cyanobacterium,Spirulina platensis strain NIES 46, were investigated using ammonium, nitrate, nitrite, or urea as the sole nitrogen source in a batch culture. Chlorophylla concentration was highest at 2,096 μg/L in the nitrate group after 10 days of cultivation, while the dry weight ofS. platensis was highest at 4.5 g/L in the ammonium group after 30 days of cultivation. The total amino acid content was highest at 174 mg/g dry weight ofS. platensis in the urea group at the end of the cultivation period, yet the amino acid patterns forS. platensis were similar for all the experimental groups. Therefore, it seemed that the growth and amino acid composition ofS. platensis varied depending on the type of nitrogen sources, while the amino acid patterns were not changed. Also, the most efficient harvesting time forS. platensis seemed to be approximately 10 days after cultivation.