Cell events i n cambium, connected with the formation and existence of a whirled cell arrangement

The history of cambium development was reconstructed on the basis of serial tangential sections of a woody tumour of <i>Picea excelsa</i> (Lam.) Lk. On the area with whirled arragement of the tracheids intrusive growth was found to occur at the lateral radial and longitudinal edges apart from its normal occurrence at the radial apical edges. Intrusive .growth at the radial lateral edge leads to forking of the fusiform initial in the cambium plane. The growth which appears on longitudinal lateral edges oocurs between the tangential walls of cells in neighbouring storeys. In such places a drastic reconstruction of the cell arrangement in neighbouring annual rings was observed. Intrusive growth. between the tangential walls leads to a whirled arrangement of tracheids.

The formation of heterogeneous storeys in the cambium on the example of Laburnum anagyroides Medik

The formation of the storeyed pattern of cambium in the first two years of cambial activity and the structure of mature cambium was examined in <i>Laburnum anagyroides</i>, a decorative shrub growing in temperate climate. A distinct storeyed pattern was observed in two-year-old stems, despite the presence of tall rays. The heterogeneous nature of storeys was revealed by the analysis of i) the number of fusiform initials forming storeys or groups of packets; and ii) the frequency of anticlinal divisions in the examined years of cambial activity and the calculation of the relative increase of the cambial circumference. The results are discussed on the basis of the recent hypothesis of intrusive growth of fusiform initial, occurring between the tangential walls of its neighbouring initial and its closest derivative.

Correlation of intrusive growth of cambial initials to rearrangement of rays in the vascular cambium

It is well documented that apical elongation of fusiform cambial initials through extension of their longitudinal edges, and their intrusion between tangential walls of the neighbouring initials and their closest derivatives cause rearrangement of fusiform cells, without increasing the cambial circumference. However, the concurrent rearrangement of rays is not fully understood. This study deals with Pinus sylvestris L., Tilia cordata Mill. and Hippophaë rhamnoides L., possessing a nonstoreyed, storeyed and double-storeyed type of cambium, respectively, and shows that the mechanism for rearrangement of ray initials is similar to the one proposed for fusiform initials, and includes multiplication of ray initials by anticlinal divisions, intrusive growth of ray initials, elimination of ray initials caused by intrusive growth of neighbouring fusiform initials, and transformation of ray initials into fusiform initials. Intrusive growth of a ray initial does not necessarily lead to the formation of a new fusiform initial, as it is dependent on the extent of the intrusive growth taken place. The extent of rearrangement of cambial cells is determined by the intensity of events occurring among the fusiform as well as ray initials. Intrusive growth of these initials does not influence the size of the cambial circumference.

Size variation in the vascular cambium and its derivatives in two Alstonia species

Two tropical tree species viz. Alstonia venenata Br. and Alstonia neriifolia Don. (Apocynaceae) were investigated to detect size variation in different elements of the cambium and its derivative tissues. Although these two species were grown under identical climatic and edaphic conditions, fusiform initial dimensions and the elements derived from them were larger in A. venenata than in A. neriifolia. Ray initials are rectangular in A. venenata but isodiametric in A. neriifolia. An appreciable increase in length was observed in the phloem and xylem ray cells when compared to the mother cells. Maximum elongation was observed in xylem fibers during differentiation from the respective fusiform initials.

Apical intrusive growth of cambial fusiform initials along the tangential walls of adjacent fusiform initials: evidence for a new concept

A new study of cambium of Pinus sylvestris L., Tilia cordata Mill. and Wisteria floribunda (Willd.) DC provides fresh clues on the cambial dynamics, rejecting the hitherto held concept that intrusive growth of the fusiform initial occurs between the radial walls of adjacent initials. It demonstrates that intrusion of the elongating initial in fact takes place along tangential walls of adjacent fusiform initials and their immediate derivatives. It also suggests a new mechanism for ‘elimination of initials’. Intrusive growth of the fusiform initial was found to begin with development of characteristic slants, representing a transitional stage of the process of transformation of periclinal walls of fusiform initial cells into radial walls, as observed in transverse sections of active cambium. The gradually progressing event comprised (a) appearance of either a triangular microspace limited by two periclinal walls of a fusiform initial and its derivative and one radial wall of another fusiform initial in the adjacent radial file, or a rhomboidal microspace enclosed by four periclinal walls of two laterally adjacent fusiform initials and their immediate derivatives, (b) intrusion of elongating tip of fusiform initial from neighbouring file into the microspace thus formed, (c) symplastic growth of the cambial cell walls in radial direction, (d) unequal periclinal divisions of fusiform initial cells while growing intrusively, and (e) unequal periclinal divisions of derivative cells not growing intrusively. Intrusive growth between periclinal walls affected rearrangement of the fusiform initials but did not add to the cambial circumference. The existing concepts of (a) intrusion of the fusiform initial between radial walls of neighbouring initials and (b) elimination of fusiform initials from cambial surface have been reassessed and redefined.

Control of fusiform initial orientation in the vascular cambium of Abies balsamea stems by indol-3-ylacetic acid

Fusiform initial reorientation, deduced from changes in tracheid arrangement, was investigated in helical bridges left after Abies balsamea stems were girdled. The first responses included an increase in the number of S- and Z-oriented anticlinal divisions without any change in their ratio, a decrease in fusiform initial length, and the formation of traumatic resin canals. Subsequently, normal tracheid differentiation resumed, the frequency of anticlinal divisions further increased, and there was an increase in the number of oriented intrusive growth events. The orientations of the anticlinal wall and of the intrusive growth were predominantly in the direction that eventually would result in the fusiform initials being aligned parallel to the bridge angle. The application of exogenous indol-3-ylacetic acid to the upper edge of a helical bridge inhibited fusiform initial reorientation, mainly by decreasing the frequency of oriented intrusive growth events. However, immediately below the application point, exogenous indol-3-ylacetic acid also induced the fusiform initials to begin temporarily to realign so as to be perpendicular to the bridge angle. The transport of a pulse of [1-14C]indol-3-ylacetic acid applied to the shoot apex was inhibited 1 day after girdling in both helical and longitudinal bridges, particularly the former, whereas it was inhibited only in helical bridges at the end of a subsequent 11-week growing period. The degree of inhibition in the helical bridges was greater the day after girdling than at the end of the growing period, during which interval fusiform initial reorientation occurred at the bottom of the bridge width. The results support the conclusion that the orientation of fusiform initials is parallel to the direction of indol-3-ylacetic acid transport.

Cyclical cell length changes in wood in relation to storied structure and interlocked grain

Interlocked grain can result from migration of a succession of left (S) and right (Z) orientational domains along the cambium. This grain pattern occurs both in species having nonstoried and those having storied cambium. In storied cambia, storey height being fixed, geometry requires that cambial fusiform initial cells undergo cyclical length changes in phase with cyclical inclination changes, e.g., a 4% length increase for a 16° inclination. We can expect the length changes to be detectable in wood if vessel member length reflects fusiform initial cell length and if the absolute amount of intrusive growth during differentiation of fibers is nearly invariant. Measurements of cells from maximum S, maximum Z, and axially aligned grain (I) sites along a radius in Entandrophragma cyclindricum and E. utile wood (storied) revealed clearly cyclical length changes in vessel members, parenchyma strands, and xylem fibers. Measurements in Nyssa sylvatica and Platanus acerifolia (nonstoried) revealed no such changes. Hence in species having storied cambium, cyclical changes in length of wood cells can accompany the slow endogenous rhythm manifested by interlocked grain. This is true because intrusive growth in these species, though extensive, does not obi iterate effects of small differences in cambial initial cell length.

The transition from nonstoried to storied cambium in Fraxinus excelsior. I. The occurrence of radial anticlinal divisions

Cell size and arrangement in layers of terminal late xylem parenchyma were interpreted as an historical record of certain types of divisions in the cambium. This allowed study of anticlinal divisions of fusiform initial cells in the cambium of Fraxinus excelsior L. Two types of anticlinal divisions were recognized: oblique (pseudotransverse) and radial (longitudinal). Type designation is, however, arbitrary, since many intermediate configurations between oblique and radial were found. With the exception of radial–lateral divisions, both oblique and radial divisions can be classified as of the left (S) or right (Z) orientational type. The types were nonrandomly distributed. A statistically significant inverse relation was found between the relative length of the anticlinal partition (RLAP) and the length of the dividing cell. Thus, short cells tend to divide radially and longer ones to divide obliquely. As an indirect result, Fraxinus cambium is a mosaic of storied and nonstoried local areas. Consequently, Fraxinus cambium, having an average (but highly variable) RLAP of just over 50%, represents an intermediate between the storied and nonstoried condition.

THE REDUCTION OF FUSIFORM CAMBIAL CELLS IN CHAMAECYPARIS AND THUJA

The loss of fusiform initials from the cambium, which is of frequent occurrence in all parts of the tree, takes place in different ways. Some cambial cells seem gradually to fail and are shortly lost from the cambium by maturation into more or less imperfect xylem or phloem elements. The majority are transversely subdivided by one or a succession of anticlinal divisions which begin near the center of the fusiform initial and usually extend to the daughter cells. The resulting segments shorten through the following periclinal divisions, some disappearing during the process of shortening and others undergoing transformation to ray initials. Nearly all new rays in the secondary body originate in this manner.