TRACHEID SIZE AND RATE OF ANTICLINAL DIVISION IN THE CAMBIUM OF CUPRESSUS

1963 ◽  
Vol 41 (8) ◽  
pp. 1187-1197 ◽  
Author(s):  
M. W. Bannan
Keyword(s):  
Arid Regions ◽  
Ring Width ◽  
Cell Length ◽  
Small Cells ◽  
Cell Multiplication ◽  
Mature Trees ◽  
Cambial Cell ◽  
Interspecific Differences ◽  
Anticlinal Division ◽  
Maximum Cell

Differences of considerable magnitude occur in the tracheid dimensions of North American species of Cupressus. In general the species in arid regions (C. arizonica, C. glabra, C. forbesii, and C. macnabiana) have smaller cells than the species in more humid localities (C. macrocarpa and C. pygmaea). Minor, in some cases questionably significant, differences in cell size are found at different sites for the same species. Noteworthy interspecific differences exist in the frequency of anticlinal (pseudotransverse) divisions involved in cambial cell multiplication. On the whole, these divisions take place at a faster rate in the species with small cells (C. arizonica, C. macnabiana) than in species with large cells (C. pygmaea). Some general relationships between rate of growth, frequency of pseudotransverse divisions, and cell length are also evident. In the stems of mature trees, maximum cell length is apparently associated with a ring width of approximately 1–1.5 mm. Reduction in ring width is accompanied by a rise in frequency of pseudotransverse division and a slight recession in cell length. Widening of the rings produces no change in the rate of anticlinal division, in relation to linear radial accretion, but cell length declines. The orientation of the partition in pseudotransverse division is usually unidirectional in neighboring cells, but reversals in tilt occur after varied intervals. Duration of the interval between reversals shows an inverse relationship with the frequency of pseudotransverse division.

Variations in cell length and frequency of anticlinal division in the vascular cambium throughout a white spruce tree

1970 ◽  
Vol 48 (7) ◽  
pp. 1363-1371 ◽  
Author(s):  
M. W. Bannan ◽  
M. Bindra
Keyword(s):  
Negative Relationship ◽  
Ring Width ◽  
Early Growth ◽  
Cell Length ◽  
Stem Length ◽  
Cell Multiplication ◽  
Great Length ◽  
Vascular Elements ◽  
Anticlinal Division ◽  
Late Growth

In the early growth of the stem, branches, and roots, the vascular elements are relatively short and the frequency of anticlinal division involved in cambial cell multiplication is high. As growth sheaths are added in the stem, length of cell increases and rate of multiplicative division declines. A similar trend occurs upward through the lower quarter to half the height of the stem. In the root system, the later growth of vertical roots is characterized by shortness of cell and high frequency of anticlinal division, and conversely, that of horizontal roots by great length of cell and low rate of anticlinal division. Although a general negative relationship exists between rate of anticlinal division and cell length throughout much of the tree, these features sometimes vary independently, and length of cell seems to be more closely related to amount of yearly radial accretion. Through the middle to late growth of the stem a negative relationship obtains between length of cell and width of annual ring, cell length maximating at a ring width of 1–2 mm. At this stage, frequency of division may fluctuate only narrowly over a considerable range of ring width. A continued decline in ring width to less than 0.5 mm, such as may occur on senescence, is accompanied by decreased cell length and accelerated anticlinal division. Length of the cell plate in anticlinal division, relative to that of the dividing cell, is greater in the early growth of the stem and branches and throughout horizontal roots than elsewhere in the tree. Most of the anticlinal divisions are pseudotransverse. The proportion of lateral divisions ranges from about 1% in the late growth of stems to 11% in horizontal roots.

TRACHEID SIZE AND ANTICLINAL DIVISIONS IN THE CAMBIUM OF LODGEPOLE PINE

1964 ◽  
Vol 42 (9) ◽  
pp. 1105-1118 ◽  
Author(s):  
M. W. Bannan
Keyword(s):  
Growth Rate ◽  
North America ◽  
Lodgepole Pine ◽  
Ring Width ◽  
Cell Length ◽  
Cell Diameter ◽  
Western North America ◽  
Breast Height ◽  
Anticlinal Division ◽  
Maximum Cell

Tracheid size and rate of anticlinal divisions in the cambium were determined for lodgepole pine at several sites in western North America ranging in elevation from 3000 to 11,000 feet. The determinations were made at breast height in trees with boles 10–20 in. in diameter. The base rate of anticlinal division was low as compared with other conifers, the average being 0.8 divisions per cm of xylem increment when ring width exceeded 1.5 mm. The frequency of division increased as annual xylem production fell below 1 mm, and rose sharply with continued reduction in ring width under 0.3 mm. Maximum cell length was associated with a ring width of 1 mm, and the length decreased in both wide and narrow rings. The tangential width of the tracheids likewise lessened as ring width fell below 1 mm, the decline becoming accentuated with continued waning of growth to less than 0.3 mm. Cell diameter differed from the overall mean by more than 1 µ at only one site after allowance was made for dissimilarities in growth rate. Cells were shortest at the two highest sites. The newly formed partition in pseudotransverse division tended to be inclined in the same direction in neighboring cells, but the percentage of divisions deviating from the preferred orientation was higher than in other conifers. Reversals in direction of tilt occurred after varied intervals, these apparently being corrective in function.

CELL LENGTH AND RATE OF ANTICLINAL DIVISION IN THE CAMBIUM OF THE SEQUOIAS

1966 ◽  
Vol 44 (2) ◽  
pp. 209-218 ◽  
Author(s):  
M. W. Bannan
Keyword(s):  
Cell Elongation ◽  
Ring Width ◽  
Cell Length ◽  
Sequoia Sempervirens ◽  
High Rate ◽  
Cambial Cell ◽  
Anatomical Features ◽  
Anticlinal Division ◽  
Cambial Cells

This report deals with the frequency of anticlinal (multiplicative) divisions in fusiform cambial cells, cell length at the time of division, and ring width as determined mostly in Sequoia sempervirens. The general relationships between these anatomical features resembled those observed in other conifers, but the frequency of anticlinal division was relatively high and the cells were long. Accordingly, the rate of cambial cell lengthening in the repetitive cycle of anticlinal division and cell elongation was found to exceed that in other conifers. Upward in the lower part of the stem the rate of anticlinal division decreased and cell length increased. A comparatively high rate of anticlinal division was maintained in the peripheral growth of large boles in both species, but cell length was greater in S. sempervirens than in S. gigantea.

CAMBIAL BEHAVIOR WITH REFERENCE TO CELL LENGTH AND RING WIDTH IN THUJA OCCIDENTALIS L.

1960 ◽  
Vol 38 (2) ◽  
pp. 177-183 ◽  
Author(s):  
M. W. Bannan
Keyword(s):  
Ring Width ◽  
Growth Patterns ◽  
Cell Length ◽  
Cell Multiplication ◽  
Radial Increment ◽  
Cambial Cell ◽  
Thuja Occidentalis ◽  
Profound Influence ◽  
Fusiform Initials ◽  
The Relationship

The relationship between ring width and length of wood cells was studied by reference to mature white cedar trees of various growth patterns. These included trees of similar diameter but diverse growth rates, trees with transition from wide to narrow rings or from narrow to wide rings in their peripheral growth, and trees with rings varying in width in different radii. Although much fluctuation occurred among individuals there was in general an inverse relationship between cell length and ring width. On the other hand, the variations in frequency of pseudotransverse divisions in the cambium, which might be expected to have a profound influence on cell length because of their involvement in cambial cell multiplication, were not obviously related to the growth rate. The frequency of pseudotransverse divisions in the fusiform initials apparently is geared neither to circumferential expansion nor to number of periclinal divisions in the cambium but rather seems to be related simply to linear radial increment.

SEQUENTIAL CHANGES IN RATE OF ANTICLINAL DIVISION, CAMBIAL CELL LENGTH, AND RING WIDTH IN THE GROWTH OF CONIFEROUS TREES

1967 ◽  
Vol 45 (8) ◽  
pp. 1359-1369 ◽  
Author(s):  
M. W. Bannan
Keyword(s):  
Cell Size ◽  
Ring Width ◽  
Cell Length ◽  
Annual Increment ◽  
Cambial Cell ◽  
Cell Dimensions ◽  
Anticlinal Division ◽  
Cambial Cells ◽  
Uniform Ring ◽  
Tree Development

The early growth of a tree is marked by a widening of the annual increments, a deceleration in rate of multiplicative (anticlinal) division of fusiform cambial cells, and an increase in cell length. Distance outward from the pith at which maximation in cell size occurs, and subsequent trends in cell dimensions, are apparently modified by rate of growth. Continuation of a uniform ring width through the middle to late years favors comparative constancy in rate of anticlinal division and cell size during that stage of tree development. Reduction in amount of annual increment in the late growth to the width optimal for cell extension, about 1 mm, induces a delayed and probably heightened maximation in cell length. Continued lessening in radial growth to an annual accretion of 0.5 mm or less, with the onset of senility, results in acceleration in rate of multiplicative division and reduction in cell length.

TRACHEID SIZE AND ANTICLINAL DIVISIONS IN THE CAMBIUM OF PSEUDOTSUGA

1964 ◽  
Vol 42 (5) ◽  
pp. 603-631 ◽  
Author(s):  
M. W. Bannan
Keyword(s):  
British Columbia ◽  
Ring Width ◽  
Cell Length ◽  
Mean Values ◽  
Second Growth ◽  
Anticlinal Division ◽  
Maximum Cell ◽  
Division Cell ◽  
Old Trees ◽  
Mother Cells

This study is concerned with certain aspects of cambial behavior and tracheid size in Douglas-fir. Geographical origin of material was Colorado, Utah, Wyoming, Washington, and several localities in British Columbia from Golden to Vancouver Island. Tracheids were smallest in all dimensions in the Colorado collections and largest in wood from coastal British Columbia. In second growth trees 35–70 years old and 10–20 in. in diameter maximum cell length was associated with a ring width of approximately 1 mm, the length declining in both wider and narrower rings. Increase in girth was accompanied by pseudotransverse divisions, these occurring mostly in fusiform cells functioning as initials, and only rarely in derived xylem mother cells. At breast height the frequency of pseudotransverse division centered about a rate of 1.3 divisions per centimeter of xylem increment when rings were 2–7 mm wide. Reduction in ring width below 1 mm occasioned a rise in frequency of anticlinal division with consequent cell shortening. In the recent growth of old trees 475–675 years old and 3–7 ft in diameter similar relationships prevailed between frequency of anticlinal division, cell length, and ring width, but mean values for rate of division and cell length differed slightly from those for young trees. Upward in the bole, to a height of 50 ft, anticlinal divisions decreased in frequency and cell length increased, but the amount of cell lengthening was greater than could be accounted for by the change in rate of division. Notable differences in rate of division occurred between neighboring trees on most sites. Although pseudotransverse divisions took place at various times during the growing season, most were aestival. Length of the partition in pseudotransverse division, although fluctuating from cell to cell, was, in general, directly related to length of the cambial cell undergoing division. The orientation of the partition in anticlinal division was usually unidirectional in neighboring cells, but occasionally an indeterminate condition prevailed. When unidirectional, reversals in direction of tilt occurred after varied intervals. Duration of the latter tended to be inversely related to the frequency of pseudotransverse division.

THE RATE OF ELONGATION OF FUSIFORM INITIALS IN THE CAMBIUM OF PINACEAE

1965 ◽  
Vol 43 (4) ◽  
pp. 429-435 ◽  
Author(s):  
M. W. Bannan
Keyword(s):  
Cell Elongation ◽  
Radial Growth ◽  
Ring Width ◽  
Total Width ◽  
Mature Trees ◽  
Cambial Cell ◽  
Anticlinal Division ◽  
Fusiform Initials ◽  
Wide Ring ◽  
Cambial Cells

This report deals with the rate and amount of elongation of fusiform initials in the cycle of cell elongation and multiplication by pseudotransverse division which accompanies circumferential expansion of the cambium. In the recent growth of mature trees the yearly amount of cambial cell elongation drops as ring width decreases, but not in proportion to the decline in ring width. The cumulative elongation through lineal series of cells, per centimeter of xylem increment, increases as ring width decreases. The amount of cell elongation during the production of several narrow rings greatly exceeds that through a single wide ring having the same total width. It seems clear that the time factor is important in the elongation of cambial cells, and to some extent the interrelated frequency of anticlinal division, particularly when radial growth declines.

THE LENGTH, TANGENTIAL DIAMETER, AND LENGTH/WIDTH RATIO OF CONIFER TRACHEIDS

1965 ◽  
Vol 43 (8) ◽  
pp. 967-984 ◽  
Author(s):  
M. W. Bannan
Keyword(s):  
Radial Growth ◽  
Lateral Roots ◽  
Height Growth ◽  
Growth Potential ◽  
Cell Length ◽  
Small Cells ◽  
Width Ratio ◽  
Mature Trees ◽  
Cell Dimensions ◽  
Length Width

In the gradual enlargement of tracheid dimensions which takes place outward from the pith in the stem and branches, growth in cell length outstrips that in tangential width so that the length/width ratio rises. Largest tracheids are usually found in the inner wood of lateral roots, where the length/width ratio is highest. Interspecific and interregional differences follow a similar pattern of relationship with respect to cell dimensions. As a rule, species or races which inhabit unfavorable sites have comparatively small cells with a low length/width ratio. Species which populate better sites and have greater growth potential are characterized by longer cells and a higher length/width ratio. Indications exist of a positive relationship between height growth, tracheid length, and the length/width ratio. On the other hand, increased radial growth in mature trees is accompanied by reduction in mean cell length and a lowering of the length/width ratio.

RAY CONTACTS AND RATE OF ANTICLINAL DIVISION IN FUSIFORM CAMBIAL CELLS OF SOME PINACEAE

1965 ◽  
Vol 43 (5) ◽  
pp. 487-508 ◽  
Author(s):  
M. W. Bannan
Keyword(s):  
Cell Multiplication ◽  
Cambial Cell ◽  
Intrinsic Factors ◽  
Ray Parenchyma ◽  
Ray Cells ◽  
Anticlinal Division ◽  
Fusiform Initials ◽  
Ray Parenchyma Cells ◽  
Cambial Cells ◽  
Resin Canals

The frequency of pseudotransverse divisions involved in cambial cell multiplication was found to be slightly higher in fusiform initials bordering on fusiform rays than in other cambial cells. The extent of difference was greater in Pinus than in Pseudotsuga or Picea. Because of the larger size of fusiform rays as compared to uniseriate rays, cambial cells adjoining the former were in contact with more ray cells per millimeter of cell length than cambial cells touching only uniseriate rays. As with the frequency of pseudotransverse division, the margin of difference in extent of ray contact was greater in Pinus than in Pseudotsuga or Picea. The evidence therefore indicates that the higher rate of pseudotransverse division in cambial cells adjoining fusiform rays was correlated with the greater area of ray contact, or more specifically, the increased contact with ray parenchyma cells. The higher rate of anticlinal division was apparently the consequence of an increase in ratio of survival of daughter initials arising in pseudotransverse division, some of the smaller newly formed initials persisting in contrast to the usual failure of similar initials situated elsewhere in the cambium. Mean height of uniseriate rays tended to increase with widening of the annual rings, but the size of fusiform rays was influenced to a much smaller degree. The frequency of fusiform rays, and horizontal resin canals, showed no consistent relationship with growth rate, but appeared to be determined by intrinsic factors.

CAMBIAL BEHAVIOR WITH REFERENCE TO CELL LENGTH AND RING WIDTH IN PICEA

1963 ◽  
Vol 41 (6) ◽  
pp. 811-822 ◽  
Author(s):  
M. W. Bannan
Keyword(s):  
Ring Width ◽  
Cell Length ◽  
Slight Reduction ◽  
Optimum Level ◽  
Annual Rings ◽  
Seasonal Growth ◽  
Uniform Rate ◽  
Maximum Cell ◽  
Cambial Cells ◽  
Picea Species

This study deals with the relationships between length of wood cells, width of annual rings, and frequency of multiplicative (pseudotransverse) divisions in fusiform cambial cells of Picea. Species investigated were P. engelmanni, P. glauca, P. mariana, and P. sitchensis. Observations were based on the recent growth of trees 10–20 in. in diameter, and in the case of P. sitchensis included trees up to 7 ft in diameter. In general, maximum cell length was associated with a ring width of 1–2 mm. Cell length declined with increase in ring width, and also decreased slightly when ring width fell below the optimum level. The frequency of pseudotransverse divisions in the cambium remained at a more or less uniform rate in trees with rings 2–8 mm wide, but rose sharply as ring width dropped below 1 mm. The slight reduction in cell length in narrow rings was apparently associated with the rise in frequency of multiplicative divisions in the cambium. Maximum cell length was observed in P. sitchensis. The shortest cells and lowest frequency of multiplicative divisions were noted in P. engelmanni. As ring width decreased a higher proportion of the pseudotransverse divisions occurred during the late seasonal growth. The orientation of the partition in pseudotransverse division tended to be unidirectional, but reversals in direction of tilt occurred after varied intervals.

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