Maderas. Ciencia y tecnología
versión On-line ISSN 0718-221X
Maderas, Cienc. tecnol. v.7 n.2 Concepción 2005
Maderas. Ciencia y tecnología 7(2):121-132
A REVIEW OF THE CONFIGURATION OF BORDERED PITS TO STIMULATE THE FLUID FLOW ♣
1Hacettepe University, Wood Products Industrial Engineering, 06532 Beytepe - Ankara, Turkey.
Corresponding author: email@example.com
Received 21.06.2005. Accepted: 08.08.2005.
As the bordered pits have generally been thought to have an influence on the refractory nature of softwoods, structural behaviour of this conducting pathways is discussed according to the published literature. Various theories on the role of bordered pits to axial flow are expounded in respect to preservative treatment. Pit aspiration is also reviewed.
Key Words: Fluid Flow, Bordered Pits, Refractory Nature, Pit Aspiration, Softwood.
It is now generally accepted that conifer wood must be regarded as a heterogeneous medium with respect to axial permeability. Erickson and Crawfort (1959) found that after air-drying, permeability to water was reduced to 1-3 % of its value for green wood. Such large changes of permeability are of great importance in wood preservation and in studies of water conduction in the living tree. In this respect, it is generally acknowledged that interfacial forces are partly responsible for many phenomena in timber drying.
According to Bolton and Koutsianitis (1980), such phenomana include the aspiration of bordered pits in coniferous species, the collapse of wood in lumber drying, the reduction of mass (liquid) flow through intercellular passageways due to the blockage of pores by gas embolisms. Components of the medium have thus been identified as tracheid lumina, bordered pits, and the porous bordered pit membranes (Bolton and Beele, 1981).
BORDERED PITS AND FLUID FLOW
For precise causes of the bordered pits to fluid flow and to clarify the reasons for the differential permeability, many anatomical studies have been carried out by several wood scientists. Literature has been well covered by such reviews as Jane (1970), Panshin and de Zeeuw (1980), Tsoumis (1991), Eaton and Hale (1993), and Langrish and Walker (1993). Some of the published literature were however listed here (see Annex).
In coniferous trees, water in sapwood is known to move longitudinally through the tracheid lumina, passing from one tracheid lumen to the next through the bordered pits. The same pathway is also used by preservative liquids when penetrating wood from a transverse surface. Both longitudinal and tangential flow paths in softwoods are predominantly by way of the bordered pits as illustrated in Fig. 1, while the horizontally aligned ray cells constitute the principal pathway for radial flow (Comstock, 1970).
The longitudinal flow of softwoods is much greater than the tangential flow due to the fact that there are fewer pitted cross walls to transverse per unit length in the longitudinal than in the tangential direction (Siau, 1984). It is therefore widely believed that in longitudinal flow through wood the greatest bulk fluid transport occurs through the bordered pits of the axial tracheids. Because tracheid lumina provide an unobstructed pathway for flow, it follows that the bordered pits will largely control the movement of fluids in conifer wood (Petty, 1970).
The number of pits per tracheid varies from 50 to 300 in earlywood with only 10 to 50 rather small bordered pits in latewood (Stamm, 1970), i.e. latewood is more permeable than earlywood in seasoned material whilst earlywood is more permeable than latewood in green wood (Petty and Preston, 1969). In green wood, water may pass from one pit aperture to the other through the pit chamber and the pit membrane pores. When wood however is dried the structure may be modified by the process of aspiration in which the torus moves across the pit chamber to seal off one of the pit apertures, thus preventing fluid flow through the pit (Fig. 2).
Fig. 1. On the left, a representation of the cellular structure of a softwood in a (TLS) tangential-longitudinal section illustrating the significance of the bordered pits in both longitudinal and tangential flow. On the right, softwood timber in the (RLS) radial-longitudinal section indicating the role of the ray cells in defining the principal pathway for radial flow (after Comstock, 1970).
As mentioned by Eaton and Hale (1993), in green softwood sapwood, longitudinal flow is favoured in the earlywood due to a number of factors, i.e. larger lumen diameters, more and larger bordered pits and larger cross-field pits. However, when dried, earlywood bordered pits aspirate due to surface tension forces and permeability is reduced. As mentioned by Tsoumis (1991), a common modification of bordered pit-pairs is the lateral displacement of the membrane. This phenomenon, called aspiration, usually occurs when sapwood is transformed into heartwood or when wood dries. Apparently, it results from high tension forces set up by menisci formed in pit apertures and in pit membrane openings through which water (sap) is moving out. In softwoods, the torus seals one of the pit apertures and, therefore, blocks the passage through the pit. Aspiration makes the wood of fir, spruce, and Douglas-fir difficult to impregnate with preservatives.
Fig. 2. The diagrammatic representation of an earlywood bordered pit in section transverse to the pit membrane. On the left, (A) Aspect in unaspirated situation, (B) Aspiration of a bordered pit during drying, i.e. the torus is pulled across the pit chamber by surface tension forces. On the right, the structure of a typical dimensions of an earlywood bordered pit: 1 tracheid wall (secondary wall), 2 middle lamella (and primary wall), 3 margo strands, 4 torus, 5 pit aperture, 6 pit chamber (after Petty, 1970).
In 1933, Phillips made what appears to be the first comprehensive study of pit aspiration. He found that drying sapwood caused a gradual increase in the number of aspirated pits with loss of moisture down to the vicinity of the fibre saturation point. At this point, virtually all the earlywood pits became aspirated, whereas about one-third of the latewood pits remained unaspirated. He described the greater tendency of latewood pits to resist aspiration to the greater rigidity of the latewood pit membrane. Liese and Bauch (1967) reported observing the same phenomena.
Accordingly, the condition of bordered pits have significant influence on permeability of softwoods. In this case, the amenability of fluid flow is usually greater in green conditions of wood than in its dried conditions. During drying, capillarity and the related surface tension of the withdrawing liquid force the pit membrane and torus against the pit opening, and hence, causing aspiration effectively.
To sum up, the mechanism of the pit aspiration could be outlined as follows:
a. the liquid transport from one tracheid to another takes place through the bordered pits quite easily in the green condition make facilitation to fluid flow: greater permeability,
b. drying can aspirate the bordered pits and cause cessation to flow: lower permeability which was due to pit membrane deflection and this phenomenon is found to be higher in earlywood than in latewood.
The most of the earlier studies were designed to observe the experimental findings to understand what extent the decrease in axial permeability is caused by aspiration and consequent reduction in the number of conducting pits, by changes in the number and size of the pit membrane pores, or by changes in the proportion of the tracheids which are conducting. Whilst decreases in the permeability of wood to liquids may be attributed to pit aspiration, little or no investigation has been made in which all are measured for the same wood specimen to study the effects of drying. In this case, Stamm (1964) described the seperate methods which exist for the estimation of all these characteristics. Petty (1969), however, developed a method which allows the pit membrane pore radius, the total number of conducting pit pores, the tracheid lumen radius and the total number of conducting tracheids to be evaluated for a single specimen from measurements of gaseous permeability at various mean pressure.
At one time it was thought that all significant resistance to flow was generated by the pit margo pores in the longitudinal tracheids. However, Petty and Puritch (1970) were able to show that at least two structural components offered resistance to flow: these components were idendified as the tracheid lumina and pit margo pores. Similarly, Smith and Banks (1970) considered that two components offered resistance to flow, but in this case the components were identified as the tracheid lumina and the entire bordered pit system. Bailey and Preston (1970) suggested that the annulus bounded by the pit border (on one side), and the torus (on the other) should offer finite and significant resistance to flow. Bolton and Petty (1975) were able to show that parts of the bordered pit system other than the pit margo pores contributed to the total resistance to flow. These authors were of the opinion that this third structural componenet was the pit aperture.
Aspiration of the bordered pits of softwoods is considered to occur because of pressure differences between adjacent tracheids which develop when the wood is drying (Jane, 1980). Liese and Bauch (1967) however showed that if water was replaced with liquids of lower surface tension, aspiration did not occur and concluded that pit membrane closed as a result of surface tension. It was further shown by Thomas and Kringstad (1971) that although surface tension was responsible for displacing the pit membrane, closing of the membrane was dependent on hydrogen bonding between the pit membrane and interior surface of the pit chamber. They concluded that, as well as having the appropriate surface tension, the evaporating liquid must be able to form hydrogen bonds possessing both hydrogen donor and acceptor properties. The liquid must also have the ability to swell wood as much as water.
The percentage of aspirated bordered pits in the transition wood of Cryptomeria japonica was determined by Nobuchi and Harada (1983). Non-aspirated pits in the sapwood rapidly became aspirated at the boundry of the transition wood whose moisture content remained high and parenchyma cells alive. Yamamoto (1982) examined the incrustation of the bordered pit membrane at the transition wood of Pinus species with UV absorbing substances (heartwood substances) and showed that it began in the innermost sapwood and preceded the decrease of moisture content. He also reported that peroxidase activity of the bordered pit membrane disappeared at the same, while its activity in ray parenchyma cells increased at the transition wood only during the growing season. Saka and Goring (1983) investigated the distribution of inorganic constituents in different morphological regions across the stem of Picea mariana. All of the elements were concentrated and localized in the tori and half-bordered pit membranes and some elements as Na, Cl, K, S and Al were only deposited in the transition wood. These results showed the screening function of bordered pit membranes and the blockage of transported water in the transition wood by the incrustation. Incrustation or aspiration of bordered pits in the innermost sapwood or outer part of the transition wood preceding the moisture decrease obviously are key factors in the explanation of the nature of heartwood formation.
Aspiration makes the most of the softwood species difficult to impregnate with preservative solutions under pressure, and hence this species became refracted, i.e. resistant to fluid flow and require a long period of treatment (EN 350, 1994 part 2). In this case, spruce sapwood is generally regarded as being very permeable to fluid before drying (Erickson, 1970) but after drying it is much less permeable (Baines and Saur, 1985) and is classed as resistant to preservative treatment (Siau, 1984). It is generally believed that the cause of loss of permeability is axial tracheid bordered pit aspiration in the earlywood where the pit margo and torus are displaced when air bubbles move past the membrane as would typically occur during drying (Petty, 1972).
According to Phillips (1933), the degree of pit aspiration in Norway spruce as in the region of 97 % while in Scots pine (which is known as a permeable species) it was in the region of 93 per cent. Although permeability is an extremely variable property of wood between the species, it is unlikely that this totally accounts for the differences in longitudinal permeability of these two softwood species.
Liese and Bauch (1967) examined the In the air-dried sapwood samples of the Pineceae, and observed that in the earlywood all bordered pits were closed, whereas in the latewood differences between species were noticed. In Abies alba, the bordered pits in the radial cell wall are mostly aspirated as in the earlywood,whereas the tangential pits, especially present in the last cell-rows of the annual rings, were open. However, in the latewood of Picea abies 20-25% of the pit membranes were found to be unaspirated. In Pinus sylvestris even up to 50% of the pits remained open. These observations provide an explanation for the differential penetrability of tracheids in earlywood and latewood of one species, and of the latewood alone between the species.
ALTERNATIVE SEASONING METHODS: SOLVENT DRYING
According to Comstock and Cote (1968), it is evident that the permeability of the wood (Abies grandis) was affected markedly by the method of drying, i.e., specimens dried after solvent exchanging water for ethanol, acetone, benzene and ethyl were examined and the invariable result was that the majority of the pits were unaspirated, which corresponds to the permeability. That the earlywood remains conducting after solvent-drying is in agreement with the findings of other workers (Erickson and Crawford, 1959; Liese and Bauch, 1967), and may be attributed to the pit aspiration of earlywood bordered pits being prevented by the absence of water at the time of drying. In Comstock and Cote′s work (1968), Red pine was reduced to about 15 percent of its original permeability and Eastern hemlock was reduced to less then 1 percent of its original permeability. There does not appear to be any trend in permeability with either surface tension or swelling of the liquids, so it seems unlikely that the higher permeability is produced by evaporation of the solvent from the wood.
In Petty and Puritc′s work (1970), the solvent dried specimens show much less variation in gas permeability than the air-dried ones. This may be explained by the presence of a varying proportion of latewood in the specimens, which does not affect the permeability of solvent-dried specimens in which virtually all of each growth ring is conducting.
Decreases in the permeability ow wood to liquids with respect to time can be attributed to the growth of gas embolisms in the flow paths. Bolton and Petty (1978) stated that, such embolisms develop in the presence of particulate gas nuclei, or where the pressure gradient in the wood is large enough to cause gas in the liquid to come out of solution. With this knowledge, many researchers have been able to obtain flow rates more nearly constant with respect to time, by subjecting their test liquids to such treatments as microfiltration, boiling or distillation, shock-cavitation, and storage under vacuum.
The bordered pits governing the longitudinal permeability of softwoods has been discussed according to the published literature. Although this review covers mainly the period from 1913 to 1987, some later studies are also included as background material.
This review will interest wood scientists concerning with wood preservation both in research institutes and at universities, and that it will suggest to some of them possible lines for research.
♣This paper was originally presented at the 36th Annual Meeting of the International Research Group on Wood Protection held in Bangalore, India, 24-28 April 2005 (Document No: IRG/WP 05-40315) but it has been revised.
BAILEY, I.W. 1913. The preservative treatment of wood, II. The structure of the pit membranes in the tracheids of conifers and their relation to the penetration of gases, liquids and finely divided solids into green and seasoned wood. Forestry Quarterly, 11: 12-20. [ Links ]
BAILEY, P.J. AND PRESTON, R.D. 1969. Some aspects of softwood permeability (I). Structural studies with Douglas-fir sapwood and heartwood. Holzforschung, 23 (4): 113-120 [ Links ]
BAILEY, P.J., PRESTON, R.D. 1970. Some aspects of softwood permeability (II). Holzforschung, 24 (2): 37-45 [ Links ]
BAINES, E.F. AND LEVY, J.F. 1979. Movement of water through wood. Journal of Institute of Wood Science, 8: 109-113. [ Links ]
BAINES, E.F. AND SAUR, J.M. 1985. Preservative treatment of spruce and other refractory wood. Proceedings, American Wood Preservation Association, 14-26. [ Links ]
BAMBER, R.K. AND FUKAZAWA, K. 1985. Sapwood and heartwood: a review. Forestry Abstracts, 46 (9): 567-580. [ Links ]
BANKS, W.B. 1970. Some factors affecting the permeability of Scots pine and Norway spruce. Journal of the Institute of Wood Science, 5 (1): 10-17. [ Links ]
BANKS, W.B. 1971. The effect of temperature and storage conditions on the phenomenon of increased sapwood permeability brought about by wet storage. Journal of Institute of Wood Science, 5 (2): 16-19. [ Links ]
BAUCH, J.; LI;ESE, W. AND SCHULTZE, R. 1972. The morphological variability of the bordered pit membranes in gymnosperms. Wood Science and Technology, 6: 165-184. [ Links ]
BOLTON, A.J. AND BEELE, P.M. 1981. The applicability of orifice flow and drag theory to the axial flow of gases through conifer wood. Wood Science and Technology, 15: 178-188. [ Links ]
BOLTON, A.J. AND KOUTSI;ANI;TI;S, G. 1980. The effect of temperature on the surface tension of sap of Thuja plicata heartwood. Wood Science and Technology, 12 (1): 3-6. [ Links ]
BOLTON, A.J. AND PETTY, J.A. 1975. Structural components influencing the permeability of ponded and unponded Sitka spruce. Journal of Microscopy, 104 (1): 33-46. [ Links ]
BOLTON, A.J. AND PETTY, J.A. 1977A. The influence of critical point and solvent exchange drying on the gas permeability of conifer sapwood. Wood Science, 9: 187-193. [ Links ]
BOLTON, A.J. AND PETTY, J.A. 1977B. Variation of susceptibility to aspiration of bordered pits in conifer wood. Journal of Experimental Botany, 28: 935-941. [ Links ]
BOLTON, A.J. AND PETTY, J.A. 1978. A model describing axial flow of liquids through conifer wood. Wood Science and Technology, 12: 37-48 [ Links ]
BRAMHALL, G. AND WI;LSON, J.W. 1971. Axial gas permeability of Douglas- fir micro-sections dried by various techniques. Wood Science, 3 (4): 223-230. [ Links ]
BUCKMAN, S.J.; SCHMI;TZ, H. AND GORTNER, R.A. 1935. A study of certain factors influencing the movement of liquids in wood. Journal of Physical Chemistry, 49 (1): 103-120. [ Links ]
BURO, A. AND BURO, E.A. 1959. Contribution to the knowledge of how liquids penetrate into pine wood. Beitrag zur Kenntnis der Eindringwege fur Flussigkeiten in Kiefernholz. Holzforschung, 13 (3): 71-77. [ Links ]
COMSTOCK, G.L. 1970. Directional permeability of softwoods. Wood and Fiber, 1(4): 283-289. [ Links ]
COMSTOCK. G. L. AND COTE, W.A. 1968. Factors affecting permeability and pit aspiration in coniferous wood. Wood Science and Technology, 2 (4): 279-291. [ Links ]
COTE, W.A. AND KRAHMER, R.L. 1962. The permeability of coniferous pits demonstrated by electron microscopy. Tappi, 45 (2): 119-122. [ Links ]
DEGROOT, R.C. AND KUSTER, T.A. 1986. SEM X-ray microanalysis of tracheid cell walls in southern yellow pine sapwood treated with water dispersible pentachlorophenol. Wood and Fiber Science, 18 (1): 58-67. [ Links ]
DUNLEAVY, J.A. AND MCQUI;RE, A.J. 1970. The effect of water storage on the cell-wall structure of Sitka spruce (Picea sitchensis) with reference to its permeability and preservation. Journal of Institute of Wood Science, 5 (2): 20-28. [ Links ]
EATON, R.A. AND HALE, M.D.C. 1993. Wood: Decay, Pests and Protection. Chapman and Hall Ltd, London (546 pp). [ Links ]
EN 350. 1994. Durability of wood and wood-based products: natural durability of solid wood. Part 2: Guide to natural durability and treatability of selected wood species of importance in Europe. British Standards Institute, London. [ Links ]
ERICKSON, H.D. 1970. Permeability of southern pine wood - A review. Wood Science, 2 (3): 149-158. [ Links ]
ERI;CKSON, H.D. AND CRAWFORD, R.J. 1959. The effects of several seasoning methods on the permeability of wood to liquids. Proceedings, American Wood Preservation Association, 55: 210-220. [ Links ]
ERI;CKSON, H.D. AND SCHMI;DT, R.N. 1969. Rupture of pit membranes during embedding procedures. Wood Science and Technology, 3 (3): 194-202. [ Links ]
ERI;CKSON, H.D.; SCHMI;TZ, H. AND GORTNER, R.A. 1938. Directional permeability of seasoned woods to water and some factors which affect it. Journal of Agricultural Research, 56 (10): 711-746. [ Links ]
GREGORY, S.C. AND PETTY, J.A. 1973. Valve action in bordered pits of conifers. Journal of Experimental Botanic, 24: 763-767. [ Links ]
HARADA, H. 1964. Further observation on the pit structure of wood. Journal of Japan Wood Research society, X: 221-225. [ Links ]
HART, C.A. AND THOMAS, R.J. 1967. Mechanism of Bordered Pit Aspiration as Caused by Capillarity. Forest Products Journal, 17 (11): 61-68 [ Links ]
HAYASHI;, S.; NI;SHI;MOTO, K. AND KI;SHI;MA, T. 1966. Study on the liquid permeability of softwoods. Wood Research, 36: 47-57. [ Links ]
JAGELS, R. 1984. PEG-isothiocyanate procedures for assessing liquid movement in wood. Pacific Regional Wood Anatomy Conference, Tsukuba, Japan. [ Links ]
JANE, F.W. 1970. The structure of wood (Ed. by Wilson, K. and White, J. B. ). Adam and Charles Black, London. [ Links ]
JOHANSSON, I. AND EDBERG, K.N. 1987. Studies on the permeability of Norway spruce. The International research Group on wood preservation, IRG/WP/2295. [ Links ]
KEI;TH, C.T. 1985. Attempts to improve penetration of waterborne preservatives in spruce and jack lumber. Forest Products Journal, 35 (11-12): 59-64. [ Links ]
KELSO, W.C.; GERTJEJANSEN, R.O. AND HOSSFELD, R.L. 1963. The effect of air blockage upon the permeability of wood to liquids. University of Minnesota, Agricultural Experiments Station, St. Paul, Technical Bulletin, 242 (40 pp). [ Links ]
LANGRI;SH, T.A.G. AND WALKER, J.C.F. 1993. Transport processes in wood. In: Primary Wood Processing, Principles and Practice (ed. by J.C.F. Walker). Chapman and Hall Ltd., London. [ Links ]
LI;ESE, W. 1965. The fine structure of bordered pits in softwoods. In: Cellular ultrastructure of woody plants. Syracuse University Press, pp. 271-290. [ Links ]
LI;ESE, W. AND BAUCH, J. 1966. Longitudinal permeability of green Silver-fir and Norway Spruce sapwood to organic solvents. Holzforschung, 20 (6): 169-74. [ Links ]
LI;ESE, W. AND BAUCH, J. 1967a. On the Closure of Bordered Pits in Conifers. Wood Science and Technology, 1 (1): 1-13 [ Links ]
LI;ESE, W. AND BAUCH, J. 1967b. On anatomical causes of the refractory behaviour of spruce and Douglas-fir. Journal of the Institute of Wood Science, 19 (1): 3-14. [ Links ]
LI;ESE, W. AND BAUCH, J. 1977. Investigations on the permeability of green sapwood of Norway spruce and Sitka spruce. Holz als Roh-und Werkstoff, 35 (7): 267-271. [ Links ]
NI;CHOLAS, D.D. AND SI;AU, J.F. 1973. Factors Influencing the Treatability of Wood. In: Wood Deterioration and Its Prevention by Preservative Treatment. Vol. 2. (ed. By D.D. Nicholas), Syracuse University Press, pp. 299-343. [ Links ]
NOBUCHI, T. AND HARADA, H. 1983. Physiological features of the “white zone” of Sugi (Cryptomeria japonica D. Don): Cytological feature and moisture content. Mokuzai Gakkaishi, 29: 824-832. [ Links ]
PALIN, M.A. AND PETTY, J.A. 1981. Permeability to water of the cell wall material of spruce heartwood. Wood Science and Technology, 15: 161-169. [ Links ]
PANSHI;N, A.J. , AND DE ZEEUW, C. 1980. Textbook of wood technology. 4th edition. McGraw-Hill Book Co., New York (728 pp). [ Links ]
PETTY, J.A. AND PRESTON, R.D. 1969. The removal of air from wood. Holzforschung, 23 (1): 9-15. [ Links ]
PETTY, J.A. 1969. Permeability and structure of the wood of Sitka spruce. Proceedings, Royal Society of London, B175 (1039): 149-166. [ Links ]
PETTY, J.A. AND PURI;TCH, G.S. 1970. The effects of drying on the structures and permeability of the woods of Abies grandis. Wood Science and Technology, 4 (2): 140-154 [ Links ]
PETTY, J.A. 1970. The relation of wood structure to preservative treatment. In: The Wood WeGrow, (ed. by The Society of Forestry Britain), pp. 29-35, Oxford University Press. [ Links ]
PETTY, J.A. 1972. The aspiration of bordered pits in conifer wood. Proceedings, Royal Society of London, B181 (1065): 395-406. [ Links ]
PETTY, J.A. 1978. Effects of solvent-exchange drying and filtration on the absorption of petroleum distillate by spruce wood. Holzforschung. 32 (2): 52-55. [ Links ]
PHI;LLI;PS, E.W.J. 1933. Movement of the pit membrane in coniferous woods, with special reference to preservative treatment. Forestry, 7: 109-120. [ Links ]
SAKA, S. AND GORI;NG, D.A.I. 1983. The distribution of inorganic constituents in black spruce wood as determinde by TEM, EDXA. Mokuzai Gakkaishi, 29: 648-656. [ Links ]
SI;AU, J.F. 1970. Pressure Impregnation of Refractory Woods. Wood Science, 3 (1): 1-7 [ Links ]
SI;AU, J.F. AND SHAW, J.S. 1971. The Treatability of Refractory Softwoods. Wood and Fiber, 3 (1): 1-12 [ Links ]
SI;AU, J.F. 1984. Transport Processes in Wood. Springer-Verlag. Berlin. p. 245. [ Links ]
SMI;TH, D.N. AND LEE, E. 1958. The longitudinal permeability of some hardwoods and softwoods. Forest Products Research, Special Report 13. HMSO, London. [ Links ]
SMI;TH, D.N. 1963. The permeability of wood to liquids and gases. Paper to FAO, 5th Conference on Wood Technology. Forest Products Laboratory, Wisconsin. [ Links ]
SMI;TH, D.N. AND BANKS, W.B. 1970. The mechanism of flow of gases through coniferous wood. Proceedings, Royal Society of London, B 177 (1047): 197-223. [ Links ]
STAMM, A.J. 1964. Wood and Cellulose Science. Ronald Press Company, New York (549 pp). [ Links ]
STAMM, A.J. 1970. Maximum effective pit pore radius of the heartwood and sapwood of six softwoods affected by drying and soaking. Wood and Fiber, 1 (4): 263-269. [ Links ]
THOMAS, R.J. AND NI;CHOLAS, D.D. 1966. Pit membrane structure in Loblolly pine as influenced by solvent exchange drying. Forest Products Journal, 16 (3): 53-56. [ Links ]
THOMAS, R.J. 1969. The ultrastructure of southern pine bordered pit membranes as revealed by specialised drying techniques. Wood and Fiber, 1 (2): 110-123. [ Links ]
THOMAS, R.J. AND KRI;NGSTAD, K.P. 1971. The role of hydrogen bonding in pit aspiration. Holzforschung, 25 (5): 143-149. [ Links ]
THOMAS, R.J. AND NI;CHOLAS, D.D. 1968. The ultrastructure of the pinoid pit in southern yellow pine. Tappi, LI: 84-88. [ Links ]
TSOUMI;S, G.T. 1991. Science and Technology of Wood: Structure, Properties, Utilisation.Van Nostrand Reinhold, New York (494 pp). [ Links ]
UNLUGI;L, H.H. 1971. Penetrability and strength of white spruce wood after ponding. Forest Products Journal, 22 (9): 92-100. [ Links ]
UNLUGI;L, H.H. 1971. Permeability of white spruce after water storage. Journal of Institute of Wood Science, 5 (6): 30-35. [ Links ]
WARDROP, A.B. AND DAVI;ES, G.W. 1961 Morphological Factors Relating to the Penetrations of Liquids into Wood. Holzforschung, 15 (5): 129-141 [ Links ]
YAMAMOTO, K. 1982. Yearly and seasonal process of maturation of ray parenchyma cells in Pinus species. Research Bulletin of Experiment Forests, Hokkaido University, 39: 245-296. [ Links ]
Title of the article from the investigation
(1913). Bailey, L.W.
The structure of the pit membranes in the tracheids of conifers and their relation to the penetration of gases, liquids and finely divided solids into green and seasoned wood.
(1933). Phillips, E.J.
Movement of the pit membrane in coniferous woods with special reference to preservative treatment .
(1935). Buckman, S.J., Schmitz, H., Gortner, R.A.
A study of certain factors influencing the movement of liquids in wood.
(1938). Erickson, H.D., Schmitz, H., Gortner, R.A.
Directional permeability of seasoned woods to water and some factors which affect it.
(1958). Smith, D.N., Lee, E.
The longitudinal permeability of some hardwoods and softwoods.
(1959). Buro, A., Buro, E.A.
Contribution to the knowledge of how liquids penetrate into pine wood.
(1961). Wardrop, A.B., Davies, G.W.
Morphological factors relating to the penetrations of liquids into wood.
(1962). Cote, W.A., Krahmer, R.L.
The permeability of coniferous pits demonstrated by electron microscopy.
(1963). Smith, D.N.
The permeability of wood to liquids and gases.
Kelso, W.C., Gertjejansen, R.O., Hossfeld, R.L.
The effect of air blockage upon the permeability of wood to liquids.
(1964). Harada, H.
Further observation on the pit structure of wood.
(1965). Liese, W.
The fine structure of bordered pits in softwoods.
(1966). Hayashi, S., Nishimoto, K., Kishima, T.
Study on the liquid permeability of softwoods.
Liese, W., Bauch, J.
Longitudinal permeability of green Silver-fir and Norway Spruce sapwood to organic solvents.
Thomas, R.J., Nicholas, D.D.
Pit membrane structure in Loblolly pine as influenced by solvent exchange drying.
(1967) Hart, C.A., Thomas, R.J.
Mechanism of bordered pit aspiration as caused by capillarity.
Liese, W., Bauch, J.
On the closure of bordered pits in conifers.
Liese, W., Bauch, J.
On anatomical causes of the refractory behaviour of spruce and Douglas-fir.
(1968). Comstock. G.L., Cote, J.R.
Factors affecting permeability and pit aspiration in coniferous wood.
Thomas, R.J., Nicholas, D.D.
The ultrastructure of the pinoid pit in southern yellow pine.
(1969). Bailey, P.J. and Preston, R.D.
Some aspects of softwood permeability (I): Structural studies with Douglas-fir sapwood and heartwood.
Erickson, H.D., Schmidt, R.N.
Rupture of pit membranes during embedding procedures.
Permeability and structure of the wood of Sitka spruce.
The ultrastructure of southern pine bordered pit membranes as revealed by specialised drying techniques
(1970). Banks, W.B.
Some factors affecting the permeability of Scots pine and Norway spruce.
Dunleavy, J.A. and McQuire, A.J.
The effect of water storage on the cell-wall structure of Sitka spruce (Picea sitchensis) with reference to its permeability and preservation.
The relation of wood structure to preservative treatment.
Petty, J.A., Puritch, G.S.
The effects of drying on the structures and permeability of the woods of Abies grandis.
Pressure impregnation of refractory woods.
Maximum effective pit pore radius of the heartwood and sapwood of six softwoods affected by drying and soaking.
(1971). Banks, W.B.
The effect of temperature and storage conditions on the phenomenon of increased sapwood permeability brought about by wet storage.
Bramhall, G., Wilson, J.W.
Axial gas permeability of Douglas- fir micro-sections dried by various techniques.
Siau, J.F., Shaw, J.S.
The treatability of refractory softwoods.
Thomas, R.J., Kringstad, K.P.
The role of hydrogen bonding in pit aspiration.
Permeability of white spruce after water storage.
Penetrability and strength of white spruce wood after ponding.
(1972). Bauch, J., Liese, W. and Schultze, R.
The morphological variability of the bordered pit membranes in gymnosperms.
The aspiration of bordered pits in conifer wood.
(1973). Gregory, S.C., Petty, J.A.
Valve action in bordered pits of conifers.
Nicholas, D.D. and Siau, J.F.
Factors influencing the treatability of wood.
(1975). Bolton, A.J., Petty, J.A.
Structural components influencing the permeability of ponded and unponded Sitka spruce.
(1977). Bolton, A.J. and Petty, J.A.
Variation of susceptibility to aspiration of bordered pits in conifer wood.
Bolton, A.J. and Petty, J.A.
The influence of critical point and solvent exchange drying on the gas permeability of conifer sapwood.
Liese, W. and Bauch, J.
Investigations on the permeability of green sapwood of Norway spruce and Sitka spruce.
(1978). Baines, E.F., and Levy, J.F.
Movement of water through wood.
Bolton, A.J., Petty, J.A.
A model describing axial flow of liquids through conifer wood.
Effects of solvent-exchange drying and filtration on the absorption of petroleum distillate by spruce wood.
(1981). Bolton, A.J., Beele, P.M.
The applicability of orifice flow and drag theory to the axial flow of gases through conifer wood.
Palin, M.A. and Petty, J.A. (1981).
Permeability to water of the cell wall material of spruce heartwood.
(1984). Jagels, R.
PEG-isothiocyanate procedures for assessing liquid movement in wood.
(1985). Keith, C.T.
Attempts to improve penetration of waterborne preservatives in spruce and jack lumber.
(1986). Degroot, R.C., Kuster, T.A.
SEM X-ray microanalysis of tracheid cell walls in southern yellow pine sapwood treated with water dispersible pentachlorophenol.
(1987). Johansson, L., Edberg, K.N.
Studies on the permeability of Norway spruce.