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1.
A system was developed to test the effects of floodwater O2concentration on ethylene evolution and stem lenticel hypertrophy,and the effects of exogenous ethylene on stem lenticel hypertrophyin mango (Mangifera indica L.) trees. Dissolved O2 concentrationsof 1–7x10–9 m3 m–3 generally resulted in hypertrophyof stem lenticels within about 6 d of flooding, whereas floodwaterO2 concentrations of 13–15 x 10–9 m3 m–3 delayedhypertrophy until about day 9. After 14d of flooding, therewere more than twice the number of hypertrophied lenticels pertree with floodwater O2 concentrations of 1–7 x 10–9m3 m–3 than with floodwater O2 concentrations of 15 x10–9 m3 m–3. Ethylene evolution from stem tissueimmediately above the floodline increased 4- to 8-fold in treesexposed to floodwater O2 concentrations of 1–2 x 10–9m3 m–3, increased 2-fold for trees exposed to floodwaterO2 concentrations of 6–7 x 10–9 m3 m–3, butremained constant with floodwater O2 concentrations of 13–15x 10–9 m3 m–3. Plants maintained in highly oxygenatedfloodwater (13–15 x 10–9 m3 m–3), and givenexogenous ethylene developed many hypertrophied lenticels, whereasplants in highly oxygenated water and not given ethylene developedfewer or nohypertrophied lenticels. These data suggest thatethylene plays a role in promotion of stem lenticel hypertrophyin flooded mango trees, and that floodwater dissolved oxygenconcentration can regulate stem lenticel hypertrophy and ethyleneevolution in this species. Key words: Flooding, hypoxia, hypertrophic cell swelling  相似文献   

2.
Seedlings of pond pine ( Pinus serotina Michx.), sand pine [ P. clausa (Engelm.) Sarg.], and loblolly pine ( P. taeda L., wet-site and drought-hardy seed sources) were grown in hydroponic solution culture using a non-circulating, continuously flowing design under anaerobic or aerobic conditions to determine whether flooding tolerance was correlated with enhanced internal root aeration. Transport of atmospheric O2 from the shoot to the root of anaerobically grown loblolly and pond pine seedlings was demonstrated via rhizosphere oxidation, using both reduced indigo-carmine solution and a polarographic, ensheathing Pt-electrode. Stem and root collar lenticels were the major sites of atmospheric O2 entry for submerged roots in these seedlings. No O2 leakage was detected from roots of aerobically grown pine seedlings. Longitudinal and radial pathways for gaseous diffusion via intercellular air spaces in the pericycle and between ray parenchyma cells, respectively, were demonstrated histo-logically in anaerobically grown loblolly and pond pines. Rhizosphere oxidation, and lenticel and aerenchyma development in roots of flood-intolerant sand pine seedlings grown in anaerobic solutions were minimal. Only 15 days of anaerobic growth conditions were necessary to increase internal root porosities of loblolly and pond pine seedlings – although not to the extent found in seedlings treated for 30 or 75 days. Histological results indicated that root tissue in the secondary stage of growth was capable of forming intercellular air spaces, demonstrating a degree of internal plasticity – at least in the more flood-tolerant loblolly and pond pine seedlings.  相似文献   

3.
Esophagi of White Leghron chick embryos, six days to hatching, were fixed in 5% glutaraldehyde in a cacodylate-sucrose buffer, postfixed in 1% OsO4, dehydrated and embedded in Araldite. Tissues were sectioned with glass knives and viewed with a Philips EM 100C. The epithelium in early stages of development is characterized by small intercellular spaces, few cell processes and few desmosomes. In contrast the differentiated epithelium contains numerous cell processes, large intercellular spaces and numerous desmosomes. Mucin appeared in the mucous glands at 17 days.  相似文献   

4.
Bramley  Helen  Hutson  John  Tyerman  Steve D. 《Plant and Soil》2003,253(1):275-286
Dieback of riparian species on floodplains has been attributed to increased soil salinisation due to raised groundwater levels, resulting from irrigation and river regulation. This is exacerbated by a reduction in flooding frequency and duration of inundation. For the Chowilla floodplain on the River Murray raised water tables have increased the amount of salts mobilised in the soil profile, causing the trees to experience salt induced water stress. For the trees to survive in the long term, salts need to be leached from the root zone.This study investigated whether floodwater infiltrates through channels created by E. largiflorens (black box) roots, flushing salts away from roots, thereby allowing the trees to increase their water uptake. Trees at different sites on the floodplain were artificially flooded, by pumping 1.5 kL of creek water into impoundments constructed around the trees. Gas exchange parameters, and pre-dawn and midday water potential were measured the day before, the day after and one week after the artificial flood and compared against trees that were not flooded. Pre-dawn and midday water potentials were also measured one month after the flood. After flooding, the trees experienced less water stress, indicated by an increase in water potential of less than 0.2 MPa, in comparison to non-flooded control trees. However, this response was not evident one month after flooding. The response to flooding did not result in increased rates of transpiration, stomatal conductance or photosynthesis, even though flooding effectively doubled the trees yearly water supply.The infiltration of floodwater in the impoundments around E. largiflorens was also compared to that of impoundments on bare ground. Floodwater infiltrated 2 – 17 times faster around trees than on adjacent bare ground, for parts of the floodplain not grazed by livestock. Tracer dye experiments indicated that bulk flow of water through pores down the profile was the reason for the enhanced infiltration. Flooding leached salts in direct vicinity of tree roots, but only leached small amounts of salts from the bulk soil.  相似文献   

5.
Nodules of the aquatic mimosoid legume Neptunia plena (L.) Benth.were always found associated with roots but not stems. Theyappeared macroscopically 10 and 20 d after inoculation on plantsgrown hydroponically and in vermiculite, respectively. The developmentof nitrogen-fixing cells occurred in a series of stages notyet reported in legume nodule formation: initial infection wasapparently intercellular and rhizobia spread between cells andthrough intercellular spaces before penetrating individual hostcells by means of infection threads. Subsequently nodule developmentwas broadly similar to that described for indeterminate papilionoidnodules. The infection threads of Neptunia and pea nodules containeda matrix with a common epitope, which was, in Neptunia, extrudedfrom the infection thread at the point of bacterial release. The central tissue contained infected and interstitial cellsand was surrounded by a three-layered cortex and a phellem.Bounding the infected region was a layer two to three cellsthick with large, unoccluded intercellular spaces. Externalto this was a layer, one or more cells thick, in which the cellwalls were interlocked, reducing the number of radially orientedintercellular spaces. The outer layer, several cells thick,contained intercellular spaces many of which were occluded.These features did not vary with growth conditions in a waywhich might influence oxygen diffusion characteristics. However,the phellem of water-cultured nodules was much more aerenchymatousthan that of vermiculite-grown nodules. Aquatic legume, Neptunia plena, nitrogen fixation, oxygen, root nodules, Rhizobium  相似文献   

6.
Microautoradiography indicated that 1-year-oldFraxinus excelsior L. stem chlorenchyma assimilated external14CO2 in mid-April, when buds were swollen, but before bud-break. The lenticel regions showed the highest amount of radioactively labeled assimilates. Labeled assimilates declined in the tangential direction with increasing distance from lenticels, suggesting that14CO2 entered the stem through the open intercellular spaces of lenticels. In the radial direction, the amount of radioactively labeled assimilates did not constantly decline with growing distance from the lenticel entrance. It was high in all lenticel phelloderm cells, which had high chlorophyll autofluorescence and very small starch grains, highest in the adjacent 4–6 rows of chlorenchyma, which had larger starch grains that increased in size towards the interior rows, and much lower in the inner cortex chlorenchyma, which had large starch grains. We suggest that the main function of the lenticel chlorenchyma (lenticel phelloderm plus 4–6 rows of adjacent cortex chlorenchyma) is the refixation of respiratory CO2 which could easily leave the stem intercellular spaces, rather than the fixation of external CO2. The lenticel chlorenchyma could reduce the loss of respiratory CO2 by its photosynthetic activity.  相似文献   

7.
Role of Ethylene in Induction of Flooding Damage in Sunflower   总被引:8,自引:0,他引:8  
The possibility that symptoms of flooding damage in plants are primarily caused by an accumulation of ethylene was investigated using pot-grown sunflower (Helianthus annuus) plants. When plants were flooded to the basal pairs of leaves, ethylene in roots and stems below the water line began to increase. This coincided with the start of hypocotyl hypertrophy and new root formation in hypocotyls, which continued for 14-16 days. There were highly significant correlations between ethylene concentration and number of roots and hypocotyl diameter. After approximately 4 days of flooding, ethylene concentrations in stems between nodes for the 1st and 3rd basal pairs of leaves started to increase, coinciding with initiation of chlorophyll breakdown and epinasty of the 2nd basal pairs of leaves. Thus, there were correlations between ethylene concentration and chlorophyll breakdown and epinasty. The lower the leaves, the more chlorophyll breakdown among 1st, 2nd, 3rd, and 4th basal pairs of leaves. The longer the flooding, the more severe the flooding damage; and even when returned to normal condition, plants flooded longer than 3 days were not able to recover from flooding damage. A gas chromatographic study revealed that Ethephon was absorbed by roots and decomposed to ethylene in the plant. Damage symptoms caused by soil application of Ethephon, such as reduced stem height, chlorophyll breakdown, epinasty of the 2nd basal pairs of leaves, and hypocotyl hypertrophy, were almost identical with those caused by soil flooding treatment. Microscopic studies revealed that radially enlarged cells and increased intercellular spaces in the cortex were the major contribution to the increased hypocotyl diameter in both flooded and Ethephon-treated plants. It is concluded that the increase in ethylene concentration in flooded plants is largely, although not exclusively, responsible for flooding damage symptoms.  相似文献   

8.
The purpose of this study was to clarify the differences of the community structure and the diversity of aquatic organisms (i) among sampling sites that the distances from inlets or outlets were different each other, and (ii) between the floodwater and the irrigation water during the crop season in a paddy field. The irrigation water was sampled from one inlet. The taxonomical groups and the number of aquatic organisms ranging in size from 30µm to 2cm in the floodwater and the irrigation water were surveyed approximately every 10days during the growth period of the rice plant. Aquatic organisms were classified mainly at the order level. Thirty-eight taxonomical groups of aquatic organisms were found in the floodwater, while 18 groups were found in the irrigation water. We were not able to find the differences of the community structure of aquatic organisms among the sites. In the floodwater, the number of taxonomical group increased and the community structure changed during the late flooding period (over 50days after the onset of flooding) at any site, while those in the irrigation water hardly changed. Although the community structure of aquatic organisms differed between the floodwater and the irrigation water throughout the flooding period, the differences became especially bigger during the late flooding period. Principal component analysis showed that three groups (Pennales, Dinoflagellida, Choreotrichida) characterized the community structure in the irrigation water. Their population densities tended to be the highest at the site near inlets and the lowest at the site far from inlets.  相似文献   

9.
Flooding is a major problem in many areas of the world and soybean is susceptible to the stress. Understanding the morphological mechanisms of flooding tolerance is important for developing flood-tolerant genotypes. We investigated secondary aerenchyma formation and function in soybean (Glycine max) seedlings grown under flooded conditions. Secondary aerenchyma, a white and spongy tissue, was formed in the hypocotyl, tap root, adventitious roots and root nodules after 3 weeks of flooding. Under irrigated conditions aerenchyma development was either absent or rare and phellem was formed in the hypocotyl, tap root, adventitious roots and root nodules. Secondary meristem partially appeared at the outer parts of the interfascicular cambium and girdled the stele, and then cells differentiated to construct secondary aerenchyma in the flooded hypocotyl. These morphological changes proceeded for 4 days after the initiation of the flooding. After 14 days of treatment, porosity exceeded 30% in flooded hypocotyl with well-developed secondary aerenchyma, while it was below 10% in hypocotyl of irrigated plants that had no aerenchyma. When Vaseline was applied to the hypocotyl of plants from a flooded treatment to prevent the entry of atmospheric oxygen into secondary aerenchyma, plant growth, especially that of roots, was sharply inhibited. Thus secondary aerenchyma might be an adaptive response to flooding.  相似文献   

10.
Growth and physiological response of woody plants to flooding have been analyzed in detail; however, relatively few studies have been oriented towards the effects of water immersion on cambial activity and wood and bark anatomy of trees that are growing in prolonged flooding conditions. These studies are important to understand the possible effects of predicted sea level rising in mangroves as a consequence of global warming. We studied five species growing in a mangrove forest, sampling three to six trees of each species, in sites that have the longest flooding period. Differences in bark appearance and phloem structure between the submerged stem portion and the portion of the stem above the water surface exist in all species. Although aerenchyma formation and stem hypertrophy are the most common events related to flooding, each type of tissue responded differently. Annona glabra L., Laguncularia racemosa (L.) Gaertn f. and Hibiscus tiliaceus L. developed rythidome. Avicennia germinans (L.) Stearn developed rythidome only in the submerged stem portion. Phyllanthus elsiae Urb., developed one periderm in both stem portions. Species that developed rythidome also developed aerenchyma between periderms and in the phellem. H. tiliaceus and P. elsiae, showed the highest values for anatomical phloem and periderm characters below water surface, while an inverse tendency was observed in A. glabra and L. racemosa, suggesting that prolonged flooding modifies vascular cambium and phellogen differently. Results indicate that sea level rising would affect distribution of the species according to their specific flooding tolerance.  相似文献   

11.
Changes in net photosynthetic rate on a leaf area basis and anatomical properties during leaf development were studied in an evergreen broad‐leaved tree, Castanopsis sieboldii and an annual herb, Phaseolus vulgaris. In C. sieboldii, surface area of mesophyll cells facing the intercellular air spaces on a leaf area basis (Smes) was already considerable at the time of full leaf area expansion (FLE). However, surface area of chloroplasts facing the intercellular air spaces on a leaf area basis (Sc), and chlorophyll and Rubisco contents on a leaf area basis increased to attain their maximal values 15–40 d after FLE. In contrast, in P. vulgaris, chloroplast number on a leaf area basis, Sc and Smes at 10 d before FLE were two to three times greater than the steady‐state levels attained at around FLE. In C. sieboldii, the internal CO2 transfer conductance (gi) slightly increased for 10 d after FLE but then decreased toward the later stages. Limitation of photosynthesis by gi was only about 10% at FLE, but then increased to about 30% at around 40 d after FLE. The large limitation after FLE by gi was probably due to the decrease in CO2 concentration in the chloroplast caused by the increases in thickness of mesophyll cell walls and in Rubisco content per chloroplast surface area. These results clearly showed that: (1) in C. sieboldii, chloroplast development proceeded more slowly than mesophyll cell expansion and continued well after FLE, whereas in P. vulgaris these processes proceeded synchronously and were completed by FLE; (2) after FLE, photosynthesis in leaves of C. sieboldii was markedly limited by gi. From these results, it is suggested that, in the evergreen broad‐leaved trees, mechanical protection of mesophyll cells has priority over the efficient CO2 transfer and quick construction of the chloroplasts.  相似文献   

12.
The objective of this study was to characterize the histodifferentiation of somatic embryogenesis obtained from leaf explants of C. arabica. Therefore, we histologically analyzed the respective stages of the process: leaf segments at 0, 4, 7, 15 and 30 days of cultivation, Type 1 primary calli (primary calli with embryogenic competence) and 2 (primary calli with no embryogenic competence), embryogenic calli, globular, torpedo and cotyledonary embryos, and mature zygotic embryos. Callus formation occurred after seven days of culture, with successive divisions of procambium cell. In this cultivation phase, it was found that Type 1 primary calli are basically formed by parenchymal cells with reduced intercellular spacing, whereas Type 2 primary calli are predominantly composed of parenchymal cells with ample intercellular spaces and embryogenic calli composed entirely of meristematic cells. After 330 days, it was evident from the differentiation of somatic embryogenesis that there was formation of globular somatic embryos, consisting of a characteristic protoderm surrounding the fundamental meristem. With the maturation of these propagules after 360 days, torpedo-stage somatic embryos arose, in which tissue polarization and early differentiation of procambial strands were verified. After 390 days, cotyledonary somatic embryos were obtained, where the onset of vessel elements differentiation was verified, a characteristic also observed in mature zygotic embryos. We concluded that somatic embryogenesis obtained from C. arabica leaves initiates from procambium cell divisions that, in the course of cultivation, produce mature somatic embryos suitable for regenerating whole plants.  相似文献   

13.
The progressive changes in measured water potential and the time to equilibration was investigated forPinus elliotii varelliotii needles in thermocouple psychrometer chambers held at 15°C, 22.5°C, and 30°C. Needles at 30°C appeared to loose semi-permeable membrane integrity after 14 hours. Needles held at 22.5°C had similar problems after 48 hours while those at 15°C were not affected. time to equilibrium was shortest at 30°C and longest at 15°C.Ethanol in the chambers had the same effect on measured water potential as did >14 hours at 30°C. Needle color, chamber odor, and pattern of measured water potential suggested an anaerobic condition in the chamber was the cause of reduced membrane integrity.Journal series paper No 7187 of the Florida, Agric. Exp. Sta.At the time the work was in progress the senior author was a graduate research assistant at University of Florida.  相似文献   

14.
Abstract

The toxicity and efficacy of using three miscible oils; cabl-2, citrole and bio-dux and a chemical insecticide (chlorpyrifos-methyl) were studied against a soft scale insect, Kilifia acuminata (Signoret) infesting mango trees. The sub-chronic toxicity of tested chemicals on white albino rats was also studied. Considering the general mean number of K. acuminata stages after 3 months post spraying, chlorpyrifos methyl and capl-2 were the most efficient compounds followed by citrol and bio-dux in descending order. Chlorpyrifos-methyl was the most effective treatment for all months after spraying, the mean reduction percentage being 94.10, 91.63 and 92.00% while it gave the highest toxic effect after three months; 90.27, 87.84 and 89.73% reduction in infestation on pre-adult, adult and gravid female stages, respectively. Results also indicate that the general mean percent reduction in K. acuminata infestation were 82.62, 86.67, 78.87 and 89.28% in treated trees by citrole, cabl-2, bio-dux and chlorpyrifos-methyl, respectively. Citrole, bio-dux, cabl-2 and chlorpyrifos-methyl caused significant increase in WBCs counts, ALT and AST activities of treated rats after 15 and 30 days from treatment comparison with control. Changes occurring in the creatinine concentration showed a significant increase in rats treated with the tested chemicals 30 days from treatment, except in the case of bio-dux oil, while there were no significant changes after 15 days in rats treated with tested oils. On the contrary, tested chemicals caused a significant decrease in RBCs count and hemogobin values after 15 and 30 days form treatment, except in rats treated with bio-dux oil. Hemoglobin content showed no significant changes over the same periods.  相似文献   

15.
A survey of distribution patterns of floodwater mosquito eggs related to environmental conditions such as moisture and plant associations was conducted by using soil samples from irrigated fields in Wroc?aw, Poland. Mosquito egg distribution was determined by repeatedly flooding the soil samples with aerated water at a temperature of 25° C. Under laboratory conditions, hatching in installments of Aedes caspius (Pallas) and Aedes vexans (Meigen) were commonly observed. The results show that ~75% of the larvae of Ae. caspius and Ae. vexans hatched after the first flooding under summer‐like conditions, whereas, following the second and third flooding, the numbers of hatched larvae were significantly lower. In our study, within one intermediate flooded field, a total of 66 plant species was identified and classified into six communities. All vegetation types were associated by varied egg densities and showed differences both in richness and Shannon‐Wiener diversity index as well as in ecological indices for moisture, soil reaction, and nutrient level. Small changes in elevation along the slope within the study area showed a large difference in the distribution of mosquito eggs. The highest average egg density was observed in zones with high occurrence of Phalaris arundinacea, usually prevalent in intermediate flooded and fertile areas. Knowledge of the indicators for the distribution of floodwater mosquito eggs in temporary breeding sites may be essential for organizing a successful, integrated mosquito control program with special regard to microbial control agents.  相似文献   

16.
Responses of Melaleuca quinquenervia seedlings to flooding   总被引:1,自引:0,他引:1  
Abstract Studies were conducted on effects of flooding for 15, 30, 60, and 90 days on morphological changes, stomatal aperture, water potential, and growth of seedlings of Melaleuca quinquenervia, a species often planted for reclamation of swamps. Flooding rapidly induced formation of many hair-like adventitious roots as well as a few thick adventitious roots that originated on the original root system. Some adventitious roots also formed on submerged portions of the stem. Melaleuca seedlings were very tolerant of flooding as shown by only slight reduction in dry weight increment of shoots after 30 days of flooding in stagnant water. Although flooding for 60 or 90 days significantly reduced dry weight increment of leaves, dry weight increment of roots was not inhibited by any flooding treatment, reflecting both degeneration of some of the original roots and compensatory growth of adventitious roots. On certain days flooding induced stomatal closure on both adaxial and abaxial leaf surfaces. Extensive production of adventitious roots and some stomatal reopening after a critical period of flooding appeared to be important factors in the flooding tolerance of Melaleuca and are consistent with its aggressiveness and vigorous growth on wet sites.  相似文献   

17.
Ultrastructural changes in tomato leaves of susceptible cv. Peto 95 and resistant cv. Ontario 7710 infected with Pseudomonas syringae pv. tomato were followed by transmission electron microscopy. Up to 48 hours from the inoculation host cells of both cultivars looked quite normal and no bacteria were visible in the intercellular spaces; bacterial cells were found only in the substomatal chambers. Afterwards, the leaf cells of cv. Peto 95 began to degenerate and bacteria invaded the intercellular spaces which seemed enlarged. After 15 days the disorganization was complete: tomato cells were plasmolyzed and the intercellular spaces were filled with bacteria. In the leaves of resistant cv. Ontario 7710 no bacteria were observed later than 48 hours and no visible modifications occurred up to 15 days after the inoculation.  相似文献   

18.
The development of the triactinomyxon stage of Myxobolus cerebralis and release of mature spores from Tubifex tubifex were shown to be temperature dependent. In the present work, the effect of temperature over a range of 5-30 degrees C on the development and release of the triactinomyxon stages of M. cerebralis was studied. Infected T. tubifex stopped releasing triactinomyxon spores 4 days after transfer from 15 degrees C to 25 degrees C or 30 degrees C. Transmission electron microscopic examinations of the tubificids held at 25 degrees C and 30 degrees C for 3 days showed that all developmental stages degenerated and transformed to electron-dense clusters between the gut epithelial cells of T. tubifex. In contrast, tubificid worms held at 5 degrees C and 10 degrees C examined at the same time were heavily infected with many early developmental stages of triactinomyxon. At 15 degrees C, the optimal temperature for development, maturing and mature stages of the parasite were evident. Infected T. tubifex transferred from 15 degrees C to 20 degrees C stopped producing triactinomyxon spores after 15 days. However, 15 days at 20 degrees C was not sufficient to destroy all developmental stages of the parasite. When the tubificid worms were returned to 15 degrees C, the one-cell stages and the binucleate-cell stages resumed normal growth. It was also demonstrated that T. tubifex cured of infection by holding at 30 degrees C for 3 weeks and shifted to 15 degrees C could be re-infected with M. cerebralis spores. The waterborne triactinomyxon spores of M. cerebralis did not appear to be as short-lived as previously reported. More than 60% of experimentally produced waterborne triactinomyxon spores survived and maintained their infectivity for rainbow trout for 15 days at water temperatures up to 15 degrees C. In natural aquatic systems, the triactinomyxon spores may survive and keep their infectivity for periods even longer than 15 days.  相似文献   

19.
Alcohol dehydrogenase (Adh) is the key enzyme in alcohol fermentation. We analyzed Adh expression in order to clarify the role of Adh of soybeans (Glycine max) to flooding stress. Proteome analysis confirmed that expression of Adh is significantly upregulated in 4-day-old soybean seedlings subjected to 2 days of flooding. Southern hybridization analysis and soybean genome database search revealed that soybean has at least 6 Adh genes. The GmAdh2 gene that responded to flooding was isolated from soybean cultivar Enrei. Adh2 expression was markedly increased 6 h after flooding and decreased 24 h after floodwater drainage. In situ hybridization and Western blot indicated that flooding strongly induces Adh2 expression in RNA and protein levels in the root apical meristem. Osmotic, cold, or drought stress did not induce expression of Adh2. These results indicate that Adh2 is a flooding-response specific soybean gene expressed in root tissue.  相似文献   

20.
Apical dieback is the predominant injury symptomatology associated with growth declines of high elevation Picea rubens Sarg, trees in North America. Histological observations of uninjured tissues and of initial injury to tissues were made to understand the mechanism of injury to this species. Observations were made of hundreds of grab samples of apparently uninjured tissues and of uninjured twigs from trees growing on mountains of the Adirondack Mt., NY; Mount Mansfield, VT; Mt. Mitchell, NC; and Clingman's Dome, TN, from March 1985 to April 1986. In the normal growth pattern of red spruce, three buds elongate from each twig terminus during spring. These buds expand into shoot increments during the growing season and three new buds will form at the tip of each of the three elongated increments. The timing of developmental events varied markedly among buds of individual trees and among trees. Bud break occurred between mid-June to the end of July. Most shoot elongation was completed and periderm formation began near the end of August or in early September. A phellogen, one cell thick, formed and a phellem layer developed from phellogen derivatives during autumn 1985. Many twig samples taken in October and November had produced only one, or at most two, phellem layers external to the phellogen during the relatively short growing season. In some samples, three or more phellem layers were present between November 1985 and March 1986. In some cases a distinct phellem was not developed at all. Usually a phelloderm one cell thick developed in autumn of the first year. Tissue necrosis occurred in twigs during their first overwintering period. Injured twigs with necrotic tissues had only one or two continuous or discontinuous phellem layers. In samples that exhibited initial injury, necrotic tissues consisted mostly of cortical cells and phloem subjacent to this meager periderm. Frequently, necrotic tissues developed initially near the bases of needles and at branch “nodes” (transition zone tissues with older twigs). In contrast, twigs of healthy appearance had two or more continuous phellem layers external to the phellogen.  相似文献   

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