Tracheids softwood

Other distinct classes of wood in a tree include the portion formed in the first 10—12 years of a tree s growth, ie, juvenile wood, and the reaction wood formed when a tree s growth is distorted by external forces. Juvenile fibers from softwoods are slightly shorter and the cell walls thinner than mature wood fibers. Reaction wood is of two types because the two classes of trees react differentiy to externally applied stresses. Tension wood forms in hardwoods and compression wood forms in softwoods. Compression wood forms on the side of the tree subjected to compression, eg, the underside of a leaning tmnk or branch. Tension wood forms on the upper or tension side. Whereas in compression wood, the tracheid cell wall is thickened until the lumen essentially disappears, in tension wood, tme fiber lumens are filled with a gel layer of hemiceUulose. 

The cells that make up the stmctural elements of wood are of various si2es and shapes and are firmly bonded together. Dry wood cells may be empty or pardy filled with deposits such as gums, resias, or other extraneous substances. Long and poiated cells, known as fibers or tracheids, vary gready ia length within a tree and from species to species. Hardwood fibers are - 1 mm long, and softwood fibers are - 3 to 8 mm. 

AG type II is most abundant in the heartwood of the genus Larix and occurs as minor, water-soluble components in softwoods. Certain tree parts of western larch (I. occidentalis) were reported to contain up to 35% AG [378]. The polysaccharide is located in the lumen of the tracheids and ray cells. Consequently, it is not a cell-wall component and, by definition, not a true hemicellulose. However, it is commonly classified as such in the field of wood and pulping research. This motivated us to include the larch AG in the review. 

In softwoods, the main cell type is the tracheid, which is often mistakenly referred to as a fibre. Tracheids constitute over 90% of the volume of most softwoods, and are the principal paper-making cells of softwoods. Their average length is usually between 2 and 4 mm, with a lengthrwidth ratio (aspect ratio) often in excess of 100 to 1, but there is a wide distribution of tracheid lengths, and it is possible for some to be as short as 1 mm and for others to be as long as 5 mm (Table 2.1). The lumen, or central cavity, is several times wider than the cell wall thickness. There is also a difference between spring wood (i.e. cells synthesised in the early part of the annual... 

Figure 2.3 A schematic representation of the structure of the primary (P) and secondary (SI, S2 and S3) cell walls of a softwood tracheid (ML = middle lamella).

Trees are classified into two major groups termed softwoods (gymnosperms) and hardwoods (angiosperms). The botanical basis for classification is whether or not the tree seed is naked as in softwoods or covered as in hardwoods. A more familiar classification, which with some exceptions is valid, is based on the retention of leaves by softwoods or the shedding of leaves by hardwoods. Thus the softwoods are often referred to as evergreen trees and hardwood as deciduous trees. The major difference with regard to wood anatomy is the presence of vessels in hardwoods. Vessels are structures composed of cells created exclusively for the conduction of water. Softwoods lack vessels but have cells termed longitudinal tracheids which perform a dual role of conduction and support. 

The anatomy of softwoods will be described first because it is less complex than hardwoods. The two main cell types which constitute softwoods are tracheids, which conduct and support, and parenchyma which store food. These two cell types can be further classified as to their orientation, that is longitudinal or transverse. Cells oriented in the longitudinal direction have the long axis of the cell oriented parallel to the vertical axis of the tree trunk whereas transversely oriented cells have their long axis at right angles to the vertical axis of the tree stem. 

Since longitudinal tracheids constitute about 90 of the volume and are therefore largely responsible for the resulting physical and chemical properties of softwoods, a detailed... 

Obviously, softwood anatomy is relatively simple as only two types of cells, longitudinal tracheids and ray parenchyma, constitute the bulk of the wood. Hardwoods have a more complex anatomy as more kinds of cells are present. The roles of conduction and support are carried out by different cells and in addition to the transverse ray parenchyma, food-storage cells oriented in the longitudinal direction are present. Parenchyma oriented longitudinally are called longitudinal or axial parenchyma. Vessel segments perform the conduction role, and fibers the support role. 

Thus, most hardwood species contain four types of cells, vessel segments, fibers, transverse and axial parenchyma, whereas most softwood species possess two types longitudinal tracheids and transverse parenchyma. 

Based on the wood anatomical descriptions presented, it is obvious that hardwoods and softwoods differ considerably from each other. For example, vessels are present in hardwoods and absent in softwoods. In hardwoods more cell types, shorter cells, more parenchyma and a more variable arrangement of cell types occur. The relative uniformity of softwood anatomy is the result of the preponderance of a single type cell, the longitudinal tracheid. 

Water conduction in a tree is made possible by pits, which are recesses in the secondary wall between adjacent cells. Two complementary pits normally occur in neighboring cells thus forming a pit pair (Fig. 1-5). Water transport between adjacent cell lumina occurs through a pit membrane which consists of a primary wall and the middle lamella. Bordered pit pairs are typical of softwood tracheids and hardwood fibers and vessels. In softwoods the pit membrane might be pressed against the pit border thus preventing water transport, since the torus is impermeable. The pits connecting tracheids, fibers, and vessels with the ray parenchyma cells are half-bordered. Simple pits without any border connect the parenchyma cells with one another. 

The wood substance in softwoods is composed of two different cells tracheids (90-95%) and ray cells (5-10%). 

Tracheids give softwoods the mechanical strength required (especially the thick-walled latewood tracheids) and provide for water transport, which... 

The average length of Scandinavian softwood tracheids (Norway spruce and Scots pine) is 2-4 mm and the width in the tangential direction is 0.02 -0.04 mm (Fig 1 -8). The thickness of earlywood and latewood tracheids is 2-4 /u,m and 4-8 /xm, respectively. 

The middle layer (S2) forms the main portion of the cell wall. Its thickness in softwood tracheids varies between 1 (earlywood) and 5 (latewood) jiim and it may thus contain 30-40 lamellae or more than 150 lamellae. The thickness naturally varies with the cell types. The microfibrillar angle (Fig. 1 -16) varies between 10° (earlywood) and 20-30° (latewood). It decreases in a regular fashion with increasing fiber length. The characteristics of the S2 layer (thickness, microfibrillar angle, etc.) have a decisive influence on the fiber stiffness as well as on other papermaking properties. 

Wood can be considered as a biological composite of hollow tubes of cellulose fibers held together by a lignin matrix gluing material. Liquids are transported up and down the trees through the tubular plumbing system. About 90 percent of the wood tissue of softwood trees consists of fiber tracheid cells for liquid conduction and support. The hardwood trees evolved after the softwoods and have specialized water conduction cells called vessels. 

The predominant cell type in softwoods is the vertically oriented (along the stem axis) longitudinal tracheid. More commonly known as fibers, these tracheids are hollow, square to rectangular in cross section, have closed and tapering ends, and are arranged so that their... 

The rays in some species also contain cells known as ray tracheids, which are similar in size to parenchyma but are dead at maturity see Figure 11). Presence and/or type of ray tracheid is sometimes a very useful diagnostic feature in identification of a particular wood genus or species. With or without ray tracheids, rays are usually several cells high, but in softwoods they are generally only one cell wide (uniseriate) (Figure 13), except in special cases where they can be up to several cells wide (multiseriate). 

The parenchyma-cell content of hardwoods is, on the average, much greater than that of softwoods. This situation is a result of the wider rays (1-50 cells) and greater ray volume of hardwoods, and also the relatively high proportion of longitudinal parenchyma (2). Additionally, the rays are all parenchyma—no ray tracheids. 

Figure 22. SEM of ray cross-field pits in softwoods as seen on the wood rcmial surface. Key A, pits to ray parenchyma (RP) in western white fir and B, pits to ray parenchyma and ray tracheids (RT) in lodgepole pine.

Many physical differences exist between latewood and early-wood see Chapter 1). In softwoods, earlywood tracheids have thin... 

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