Stem phototropism of trees: a possible significant factor in determining stem inclination on forest slopes

BACKGROUND AND AIMS: The main stems of trees on forest slopes incline down the slope to various extents that are characteristic of the species. The inclination has been explained as an active response to a horizontally asymmetrical light environment, but the contributing physiological mechanisms are unknown. The present study tested the hypothesis that stem phototropism, gravitropism, or a combination of the two determines the inclination of tree stems on forest slopes. METHODS: Cryptomeria japonica, Pinus densiflora, Quercus myrsinaefolia and Q. serrata were studied. Measurements were made of stem inclination of mature trees on forest slopes in uniform plantations of each species, and changes in stem inclination of potted seedlings in response to illumination treatments (unilateral or overhead) and inclination treatments (artificially inclined or erect). Indices of phototropic and gravitropic responsiveness were evaluated for each species, calculated from the change in stem inclination in response to artificial inclination with unilateral or overhead illumination. KEY RESULTS: Stem inclination on forest slopes varied significantly among species: Q. serrata inclined most in the down-slope direction, C. japonica inclined the least, and P. densiflora and Q. myrsinaefolia were intermediate. The change in stem inclination of seedlings in each treatment varied significantly among species. One-year-old stems of Q. serrata and 2-year-old stems of Q. myrsinaefolia bent toward the light source. Interspecific variation in the change in stem inclination in response to the unilateral illumination or that in the index of phototropic responsiveness was strongly correlated with the variation in stem inclination on forest slopes. CONCLUSIONS: The orientation of woody stems that have finished elongation can be actively controlled by phototropism. Interspecific variation in phototropic responsiveness of trees is a possible significant determinant of interspecific variation in stem inclination on forest slopes.     

Phototropic bending of non-elongating and radially growing woody stems results from asymmetrical xylem formation

Active phototropic bending of non-elongating and radially growing portion of stems (woody stems) has not been previously documented, whereas negative gravitropic bending is well known. We found phototropic bending in woody stems and searched for the underlying mechanism. We inclined 1-year-old Quercus crispula Blume seedlings and unilaterally illuminated them from a horizontal direction perpendicular to ('normal' illumination) or parallel to ('parallel' illumination) the inclination azimuth. With normal illumination, active phototropic bending and xylem formation could be evaluated separately from the negative gravitropic response and vertical deflection resulting from the weight of the seedlings. One-year-old stems with normal illumination bent significantly, with asymmetrical xylem formation towards the illuminated upper surface and side of the stem, whereas those with parallel illumination showed non-significant lateral bending, with asymmetrical xylem formation only on the upper side. A mechanical model was built on the assumption that a bending moment resulted from the asymmetrical xylem formation during phototropic bending of the woody stems. The model fitted the relationship between the observed spatial distributions of the xylem and the observed lateral bending, and thus supported the hypothesis that phototropic bending of woody stems results from asymmetrical xylem formation, as such occurs during gravitropism.     

Ethylene in relation to compression wood formation in Abies balsamea shoots

 The terminal (1-year-old) shoot of dormant, 2-year-old balsam fir [Abies balsamea (L.) Mill.] seedlings was either left vertically oriented or tilted to an angle of 60° from the vertical (tilting experiment), or was ringed with N-1-naphthylphthalamic acid (NPA), an inhibitor of indole-3-acetic acid transport, at a concentration of 0, 1 or 10 mg g⁻¹ lanolin (NPA experiment). After 6 weeks of growth, ethylene evolution from the cambial region was measured by gas chromatography – flame ionization detection, and tracheid production and compression wood formation were determined by microscopy. In vertical seedlings of the tilting experiment and in 0 mg g⁻¹-treated seedlings of the NPA experiment, compression wood was not formed and neither ethylene evolution nor tracheid production varied longitudinally or circumferentially within the stem. Tilting induced compression wood formation and increased ethylene evolution and tracheid production on the lower side of the stem, while decreasing tracheid production on the upper side. Compression wood formation was induced and tracheid production and ethylene evolution were stimulated at and above the point where 1 or 10 mg NPA g⁻¹ was applied, whereas below this point compression wood was not formed and tracheid production was inhibited. In both tilting and NPA experiments, there was a positive correlation between ethylene evolution and tracheid production when data from all seedlings were analyzed, but not when data from seedlings forming compression wood were excluded. The results indicate that cambial region ethylene evolution is enhanced when compression wood is being formed, and that the enhancement is related to compression wood formation per se rather than the associated increase in tracheid production. Received: 19 February 1998 / Accepted: 26 October 1998     

龟纹瓢虫的捕食胁迫作用对连续三代果蝇发育与繁殖的影响

捕食性天敌不仅直接捕食害虫,而且还能改变害虫的生长发育、繁殖,间接地影响害虫的适合度。以常见模式昆虫黑腹果蝇Drosophila melanogaster Meigen及捕食性天敌龟蚊瓢虫为研究系统,研究龟纹瓢虫Propylea japonica Thunberg连续多代直接捕食胁迫、一代捕食胁迫后下一代不捕食胁迫,对果蝇生长发育、繁殖和适合度的影响。结果表明:如果持续地释放瓢虫对果蝇的捕食胁迫作用,显著缩短了第1代、第2代果蝇的幼虫的发育历期,增加了第1代、第2代果蝇的繁殖量。但在第3代的影响却是延长了果蝇的幼虫的发育历期,降低第3代果蝇的繁殖量。如果在上一代胁迫作用得到释放后,下一代不再有瓢虫胁迫,那么连续2代胁迫后去掉胁迫的果蝇比只胁迫一代后去掉胁迫的果蝇幼虫的发育历期明显延长,繁殖量明显下降,后代性比也明显下降。与连续胁迫条件下的处理相比,成虫的寿命明显延长;对果蝇幼虫的发育历期、繁殖量则没有显著影响。结果表明,龟蚊瓢虫的干扰胁迫作用,可以间接的增加果蝇的适合度,促进果蝇种群的增长,使果蝇种群自我维持、调节到相应的水平。     

Annual growth rings, rainfall-dependent growth and long-term growth patterns of tropical trees from the Caparo Forest Reserve in Venezuela

1 Tree-ring analyses and dendrometer measurements were carried out on 37 tree species in a semi-deciduous forest of the Reserva Forestal de Caparo, Venezuela, where the mean annual rainfall is about 1700 mm and there is a dry season from December to March. The main purposes of the investigation were to show the seasonality of cambial growth, and the connection between precipitation patterns and tree-ring curves. Long-term rates of wood increment were also estimated.
2 Cambial markings in consecutive years showed that annual rings were formed by many species.
3 The distinctiveness of growth zones was usually greater in deciduous species than in evergreen species, although not all deciduous species had distinct rings.
4 Dendrometer measurements showed that the annual growth rhythm was related to precipitation patterns. Evergreen species tended to show only a short interruption of wood growth (during the later part of the dry season), whereas deciduous species stopped growth completely at the end of the rainy season.
5 For deciduous species, regression analyses showed close relations between tree-ring width and the sum of precipitation outside the rainy seasons (i.e. November to April). Evergreen species reacted to the total annual amount of precipitation.
6 Variation in longest available ring chronology (for Terminalia guianensis ) showed little correlation with the El Niño–Southern Oscillation effect.
7 On average trees from natural forests showed relatively constant growth over the entire life span. Plantation trees grew fast up to an age of 15–20 years, but annual increments then decreased to values seen in natural forest trees.     

Discussion of development processes in insect‐fungus association derived from the shaggy parasol fruiting on the nests of hairy wood ants

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