Effects of environmental factors on development of wood

This research tested the hypothesis that environmental factors (light, water, and nutrient levels) affect wood development. Specimens were placed in treatments of low, medium, or high levels of light, water, nitrogen, or phosphorus for one year. Control plants received medium levels of all factors, while experimental plants received medium levels of all factors except the experimental factor; for example, "high light" treatment consisted of high light but medium levels of water, nitrogen, and phosphorus. Some character changes seen in Cereus peruvianus were a reduction in mean vessel diameter and shoot elongation as a result of low nitrogen and low phosphorus treatments and a reduction in mean vessel density due to low light; high water induced broader vessels and greater shoot elongation. In Cereus tetragonus, low water treatment caused a reduction in mean vessel diameter, and high nitrogen decreased the amount of wood produced. Whereas all characters studied showed a significant correlation with at least one treatment in one species, few characters responded similarly between species. Estimated specific conductivity of wood could be altered by treatments affecting either vessel density or vessel diameter strongly or by treatments affecting both diameter and density weakly. Under the conditions tested, wood structure was stable but estimated conducting capacity was more flexible.     

CYTOKININ-ELICITED FORMATION OF THE PITH-RIB MERISTEM AND OTHER EFFECTS OF GROWTH REGULATORS ON THE MORPHOGENESIS OF ECHINOCEREUS (CACTACEAE) SEEDLING SHOOT APICAL MERISTEMS

Shoot apical meristems of Echinocereus engelmannii have only a tunica-corpus organization at germination, but the corpus rapidly develops central mother cells, a peripheral zone and a pith-rib meristem. The manner in which nutrition, darkness and various growth regulators at several concentrations and in several combinations affect the development of zonation was examined by growing derooted seedlings on agar which contained the nutrients or growth regulators. Benzylaminopurine was able to elicit the formation of the pith-rib meristem in an otherwise non-zonate corpus. Also, the rate of leaf initiation was greatly increased. Gibberellic acid severely inhibited the formation of corpus zones but had little effect on leaf initiation. Indoleacetic acid had no effects other than mild inhibition of zonation and a slight retardation of leaf initiation. Abscisic acid was strongly inhibitory. Sucrose only slightly increased the rate of leaf formation and did not affect apex size or zonation. To more closely examine the cytokinin-induced effects on the apical meristem, several growth regulators were applied in combination with the most effective concentration of cytokinin. Certain combinations were able to interfere with several of the cytokinin-induced responses, while other cytokinin-induced responses occurred even in the presence of high concentrations of these other growth regulators. Leaf initiation and meristem morphogenesis appeared to be remarkably stable and insensitive to the presence of most hormones except cytokinin and gibberellin.     

THE STRUCTURE AND DEVELOPMENT OF AN UNUSUAL TYPE OF ARTICULATED LATICIFER IN MAMMILLARIA (CACTACEAE)

Although the laticifers of several species of Mammillaria can technically be classified as being of the articulated type, they differ significantly from all other reported articulated laticifers. They are derived from cells which differentiate only in older tissues, never in meristematic or young regions. The development involves the complete lysis of masses of cells, not just the perforation or resorption of the end walls in a single file of cells. At maturity, the laticifer lumen is lined with a one-to-several layered epithelium which may be quite thick. The laticifers increase in diameter with age, apparently by the lysis of the inner epithelial cells. Laticifers occur in the pith, cortex and tubercles of the vegetative body but were not observed in the roots, flower parts or in seedlings up to eight months old. Seven species were studied, all of which have “milky sap.” and the laticifers of each were virtually identical to the laticifers of the others.     

RELATIONSHIPS AMONG SHOOT APICAL MERISTEM ONTOGENIC FEATURES IN TRICHOCEREUS PACHANOI AND MELOCACTUS MATANZANUS (CACTACEAE)

Various morphometric parameters of shoot apical meristems were measured during the ontogeny of Melocactus matanzanus and Trichocereus pachanoi. While some ontogenic features varied dramatically (i.e., apical surface curvature, dK/dl, varied by 0.15–0.17, while the apical volume increased from 10⁵ μm³ to 10⁷ or 10⁸ μm³), the percent volume of the central-mother-cell zone (CMCZ), pith-rib meristem (PRM), and the peripheral zone (PZ) showed remarkable intraspecific constancy: CMCZ = 9.2%, PRM = 10.4%, PZ = 80% for M. matanzanus, and CMCZ = 10%, PRM = 24.5%, PZ = 65% for T. pachanoi. Seedlings (2 plastochrons old) showed distinctly different percent volumes from older forms: CMCZ = 7.8%, PRM = 5.9%, PZ = 86% for M. matanzanus, and CMCZ = 8.9%, PRM = 13%, PZ = 77% for T. pachanoi. Geometric expressions of cell patterns within zones of the apex remained relatively constant throughout development. On the basis of intra- and interspecific comparisons of the percent volumes among 23 taxa and the chronology in the appearance of zones within the apex, it is concluded that while there are no apparent biological reasons for the actual percent volume values computed from our sample set, the constancy in zones reflects a high degree of homeostasis which is achieved early in ontogeny.     

A MORPHOMETRY STUDY OF THE ULTRASTRUCTURE OF ECHINOCEREUS ENGELMANNII (CACTACEAE). II. THE MATURE,ZONATE SHOOT APICAL MERISTEM

The shoot apical meristems of adult Echinocereus engelmannii plants are zonate and have a tunica, central mother cells, a peripheral zone, and a pith-rib meristem. An ultrastructural, stereological study showed that each zone has its own distinct ultrastructure, but that the differences between the zones are quite small, both on a protoplasmic basis and on a cytoplasmic basis. Furthermore, the ultrastructure present in the adult apices differed only slightly from that which had been found in seedling apices, demonstrating a long-term stability of structure. The standard deviations found in the sample were small, indicating little variability from one plant to the next and suggesting that there are little or no cyclic changes during the plastochron or a 24-hr photoperiod. The ultrastructures found in the shoot apical meristems differed significantly and markedly from mature tissues of the same plants.     

Evolution of extreme xeromorphic characters in wood: a study of nine evolutionary lines in Cactaceae

Vessels of xeric-adapted woods have been predicted to be narrower than those of mesic-adapted woods, to occur at higher densities, to occur in larger clusters, and to have a greater percentage of them in clusters. These predictions were tested by comparing wood structure of several evolutionary lines of xeric-adapted cacti to that of mesic-adapted Pereskia, which probably resembles the ancestral cacti. Although derived cacti occur in habitats with water stress ranging from mild (rain forests) to severe (open deserts with little vegetation other than cacti), as long as plants retain wood with an ordinary fibrous matrix, wood characters are remarkably uniform and not correlated with habitat aridity. However, in several evolutionary lines, novel wood types occur with characters that fulfill the predictions for xeric-adapted woods listed above. However, conductive area (fraction of wood transverse-sectional area occupied by conduits) and estimated specific conductance (conductance per square millimetre) are correlated with shoot height (the need for mechanical support from xylary fibers) rather than with habitat aridity: tall plants transport water through relatively few, wide vessels, permitting much of the wood volume to consist of fibers. Small plants with little wood use large numbers of narrow vessels rather than small numbers of wide ones, thereby achieving conductive safety.     

A MORPHOMETRIC STUDY OF THE ULTRASTRUCTURE OF ECHINOCEREUS ENGELMANNII (CACTACEAE). VI. THE INDIVIDUALIZED ULTRASTRUCTURES OF DIVERSE TYPES OF MERISTEMS

The structures of spine basal meristem cells were examined by stereological morphometric techniques and found to be different not only from shoot apical meristem cells but also from spine primordium cells in the relative volumes occupied by cellular organelles. Nineteen types of meristematic cells have been studied in E. engelmannii, and all are distinct ultrastructurally. This suggests that whereas they all appear to be primarily involved in mitosis and cytokinesis, there must be other important aspects to their physiology; possibly these cells have already begun differentiation toward their mature states even while they are parts of active meristems.     

HORMONAL CONTROL OF ORGANOGENESIS IN OPUNTIA POLYACANTHA (CACTACEAE)

An axillary bud (areole) of Opuntia polyacantha is composed of an extremely short axis which bears highly modified leaves (spines). After producing the spines, the bud apical meristem becomes quiescent. When the axillary bud is excised and cultured with benzylaminopurine (BAP), the apical meristem increases in complexity and produces leaves on an elongated axis. Gibberellic acid (GA) induces spine production with no elongation of the axis. Naphthaleneacetic acid (NAA) causes no structural change in the meristem but induces root formation in adjacent tissue. The initiation of roots, spines, or leaves is visible within three days. Explants on medium lacking hormones show no structural change, but by the tenth day are insensitive to BAP or NAA. These aged explants can be made to respond to BAP or NAA by merely rewounding them at the time of hormone application. With concentrations previously found to be optimal for single hormone responses, NAA in combination with BAP prevents shoot formation, or in combination with GA prevents spine production. When all three hormones are combined, spines are produced, indicating that GA is active morphologically while BAP and NAA counteract each other. If the level of NAA is decreased, BAP counteracts the NAA and also overrides the GA and leafy shoots are produced. By varying the proportions of BAP and GA, in the absence of NAA, four types of lateral appendage can be produced: leaves, mildly altered leaves, leaf-spine transition forms, and spines.     

A MORPHOMETRIC STUDY OF THE ULTRASTRUCTURE OF ECHINOCEREUS ENGELMANNII (CACTACEAE). III. SUBAPICAL AND MATURE TISSUES

A stereological, morphometric study of the ultrastructure of subapical cells, xylem parenchyma, and cortical chlorenchyma of Echinocereus engelmannii shows that each of these cell types has a unique organellar composition. The cells of any of these tissues are unique not only in vacuolation (which is visible at the light microscope level), but also in the nucleus/cytoplasm ratio and the concentrations of mitochondria, chloroplasts, and dictyosomes. Furthermore, the differences between each of these cell types were large and statistically significant. It had previously been found that the cells of each zone of the shoot apical meristems of E. engelmannii are different from those of the other zones, but the present study suggests that, considering the large ranges of structure possible in the nonapical cells of this species, the apical meristem variation should be considered as only a minor difference and that the meristem zones are really quite similar to each other.     

The effect of the pine woolly aphid (Pineus pini) on survival,growth and natural selection in Scots pine (Pinus sylvestris) in Iceland

1. Scots pine Pinus sylvestris was originally introduced to Iceland in the beginning of the 20th Century. Extensive plantings started in the late 1940s and, in total, 2–3 million Scots pine seedlings were planted, mainly originating from two counties in northern Norway. Part of this plant material was imported as seedlings.
2. Pine woolly aphid Pineus pini was introduced to Iceland before 1940, most likely on imported seedlings in 1937.
3. High mortality of Scots pine, concurrent with high infestation of the pine woolly aphid, was observed in Iceland during the late 1950s and 1960s and planting was discontinued.
4. Provenance trials with Scots pine were established in Iceland in 2004–2006. They consisted of 15 provenances from Norway, four from Finland, four from Scotland, one from Russia, one from the Austrian Alps and three first generation Scots pine provenances from Iceland, collected from survivors of the epidemic in the 1950s and 1960s. In total, there were 28 provenances.
5. The Icelandic provenances had significantly lower P. pini infestation than all the provenances of non‐Icelandic origin, which indicates that natural selection in Scots pine in Iceland has occurred in favour of individuals less susceptible to P. pini.