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京都大学 / 新制・課程博士 / 博士(農学) / 甲第23525号 / 農博第2472号 / 新制||農||1087(附属図書館) / 学位論文||R3||N5356(農学部図書室) / 京都大学大学院農学研究科森林科学専攻 / (主査)教授 梅村 研二, 教授 矢野 浩之, 教授 仲村 匡司 / 学位規則第4条第1項該当 / Doctor of Agricultural Science / Kyoto University / DFAM

Pinus radiata is the dominant species of the plantations forests in New Zealand. The forest industry in New Zealand is heavily dependant on it. However, Pinus radiata can develop wood quality flaw called 'intra-ring checking'. The checks or splits appear in wood during kiln drying and usually affect the earlywood region of the wood. It lowers value of appearance grade timber leading to huge economic loses for the forest industry. This thesis presents a study that was undertaken as a part of ongoing collaborative work that is being carried out to understand wood quality issues in Pinus radiata, with a vision of improving its wood quality. This study was a part of that effort and was conducted with an aim to gain an insight into intra-ring checking, and the process of xylogenesis in Pinus radiata. The investigations for this study were carried out in two steps. The first step was to understand intra-ring checking. The location of intra-ring checking was determined by observing the checks using various microscopy techniques. Scanning electron microscopy confirmed that checking was as an intercell failure that usually occurs at the cm1/S1 boundary. A comparative study was also conducted to see if the checked wood had some inherent properties that made it more susceptible to checking. It was found that checking could be influenced by tracheid geometry and cell wall thickness. If the wood had large tracheids with thin walls, it was more likely to develop checks during drying. Lignin distribution in the cell wall layers was also seen to play an important role in checking. Lower lignin levels and disruption in the pattern of lignification of the cell wall layers increased the tendency of the wood to develop checks. Similarly, it the tracheids have larger pits then their tendency to check increases. Structural features that disrupt the uniformity of the interlocking pattern of the tracheid such as rays and resin canals could also play a role in checking. Checked wood tends to have more surface area occupied by ray tissue. However, resin canals do not seem to be directly involved in checking, though their arrangement could indicate disturbances during xylogenesis. The second step was to understand the process of xylogenesis in Pinus radiata especially with respect to the influence of auxin and boron on it. Nutrient and organ culture methods were manipulated and successfully used to study xylogenesis. An exhaustive comparative study was carried out to observe and measure selected wood properties. Microscopy and image analysis revealed that auxin and boron changes in the medium led to the alterations in the cell division, expansion and lignification. However, the analysis of the measurements and the observations displayed complex 'between-tree' and 'within-culture variations'. Clear trends did not emerge from the analysis hence, a confident conclusion on the association between auxin, boron and lignification could not be drawn from this organ culture study. The study has added to the knowledge about checking and wood properties associated with it. A new tool of organ culture had been established that can hlep future research on the process of xylogenesis in Pinus radiata.

Pinus radiata is the dominant species of the plantations forests in New Zealand. The forest industry in New Zealand is heavily dependant on it. However, Pinus radiata can develop wood quality flaw called 'intra-ring checking'. The checks or splits appear in wood during kiln drying and usually affect the earlywood region of the wood. It lowers value of appearance grade timber leading to huge economic loses for the forest industry. This thesis presents a study that was undertaken as a part of ongoing collaborative work that is being carried out to understand wood quality issues in Pinus radiata, with a vision of improving its wood quality. This study was a part of that effort and was conducted with an aim to gain an insight into intra-ring checking, and the process of xylogenesis in Pinus radiata. The investigations for this study were carried out in two steps. The first step was to understand intra-ring checking. The location of intra-ring checking was determined by observing the checks using various microscopy techniques. Scanning electron microscopy confirmed that checking was as an intercell failure that usually occurs at the cm1/S1 boundary. A comparative study was also conducted to see if the checked wood had some inherent properties that made it more susceptible to checking. It was found that checking could be influenced by tracheid geometry and cell wall thickness. If the wood had large tracheids with thin walls, it was more likely to develop checks during drying. Lignin distribution in the cell wall layers was also seen to play an important role in checking. Lower lignin levels and disruption in the pattern of lignification of the cell wall layers increased the tendency of the wood to develop checks. Similarly, it the tracheids have larger pits then their tendency to check increases. Structural features that disrupt the uniformity of the interlocking pattern of the tracheid such as rays and resin canals could also play a role in checking. Checked wood tends to have more surface area occupied by ray tissue. However, resin canals do not seem to be directly involved in checking, though their arrangement could indicate disturbances during xylogenesis. The second step was to understand the process of xylogenesis in Pinus radiata especially with respect to the influence of auxin and boron on it. Nutrient and organ culture methods were manipulated and successfully used to study xylogenesis. An exhaustive comparative study was carried out to observe and measure selected wood properties. Microscopy and image analysis revealed that auxin and boron changes in the medium led to the alterations in the cell division, expansion and lignification. However, the analysis of the measurements and the observations displayed complex 'between-tree' and 'within-culture variations'. Clear trends did not emerge from the analysis hence, a confident conclusion on the association between auxin, boron and lignification could not be drawn from this organ culture study. The study has added to the knowledge about checking and wood properties associated with it. A new tool of organ culture had been established that can hlep future research on the process of xylogenesis in Pinus radiata.

Exploring how drought influences growth, performance, and survival in different species is crucial to understanding the impacts of climate change on forest ecosystems. Here, we investigate the responses of two co-occurring pines (Pinus nigra and Pinus sylvestris) to interannual drought in east-central Spain by dendrochronological and wood anatomical features integrated with isotopic ratios of carbon (delta C-13) and oxygen (delta O-18) in tree rings. Our results showed that drought induces both species to allocate less carbon to build tracheid cell-walls but increases tracheid lumen diameters, particularly in the transition wood between early and latewood, potentially maximizing hydraulic conductivity but reducing resistance to embolism at a critical phase during the growing season. The thicker cell-wall-to-lumen ratio in P. nigra could imply that its xylem may be more resistant to bending stress and drought-induced cavitation than P. sylvestris. In contrast, the higher intrinsic water-use efficiency (iWUE) in P. sylvestris suggests that it relies more on a water-saving strategy. Our results suggest that narrower cell-walls and reduced growth under drought are not necessarily linked to increased iWUE. At our site P. nigra showed a higher growth plasticity, grew faster and was more competitive than P. sylvestris. In the long term, these sustained differences in iWUE and anatomical characters could affect forest species performance and composition, particularly under increased drought stress.

To evaluate the compression wood response in eastern white pine (Pinus strobus L.) following a severe ice storm in 1994, 47 trees were felled in 2007 and cross-sectional samples were collected at 0.5 (±0.2) m stem height. The disks were sanded and digitally scanned, and the cross-sectional area (mm2) of compression wood within each tree-ring was quantified using image analysis software. Topographic data (slope, aspect, and elevation) were also recorded for each P. strobus tree, along with a modified competition index. Wood anatomical features were also quantified in the three years before and after the storm along a tree diameter gradient. Although tree age was relatively constant in this stand, tree size was influenced by topographic position; larger trees grew in the valley while smaller trees were found growing in thin soils at the mid-slope position. When the cohort was about 25 years old, ice deposition caused a heterogeneous compression wood response which was highly related to tree size. In the thirteen years following the ice storm, the 6 – 9 cm (2007) diameter class formed significantly more compression wood area than any other, followed by the 10 – 13 cm (2007) diameter class. The tree diameter range that formed the most post-storm compression wood was 4 – 8 cm at the time of the storm, suggesting that this diameter range was most affected by 8.5 cm of ice loading in P. strobus. Trees > 18 cm in 1994 did not form any compression wood after the storm, but many experienced a growth release to fill canopy gaps. Topographic variables did not influence compression wood formation directly, but only one plot was sampled so these results are tenuous. However, topography did influence tree size which was the most important predictor in compression wood. There was no relationship between compression wood area and competition index. Due to compression wood formation after the ice storm, cell wall thickness and cell circularity were significantly higher in the 1994 tree-ring than in other rings examined (1991 – 1993, 1995, and 1996). Tracheid and lumen diameters were significantly smaller in compression wood cells (30.5 and 19.5 μm, respectively) than in normal wood (36.8 and 28.4 μm, respectively); opposite wood cells were intermediate in size (32.4 and 24.4 μm, respectively). Due to small tracheid size, compression wood contained significantly more cells mm⁻¹ (33) than normal wood (27), but no significant differences in cell wall area. Therefore, cumulative cell wall area occupied 47% of the cross-section in compression wood tissue on average, compared to 31% in normal wood. Dispersing tree weight across a greater surface area may help compression wood to prop up a bent tree, but reduced lumen area may also impact hydraulic conductivity in the stem. / Master of Science

Anacampserotaceae pertence à ordem Caryophyllales, subordem Portulacineae, atualmente constituída por três gêneros (Anacampseros L., Grahamia Gill ex Hook e Talinopsis A. Gray) antes pertencentes à família Portulacaceae. Os representantes de Anacampserotaceae apresentam estruturas axilares lignificadas muito características, como tricomas, escamas (scales) e cerdas (bristles) e também apresentam células do tipo WBT que ocorre somente em mais outras duas famílias de Caryophyllales, Aizoaceae e Cactaceae. Os caracteres axilares são muito informativos para o clado ACPT (Anacampserotaceae, Cactaceae, Portulacaceae e Talinaceae) e vêm revelando importantes dados aos estudos filogenéticos da subordem Portulacineae, no entanto, há uma controvérsia quanto a origem ontogenética das estruturas axilares de Anacampserotaceae, o que pode levar a interpretações equivocadas das relações filogenéticas do grupo. Diante disso, este trabalho realizou um estudo ontogenético das estruturas presentes na região da axila foliar de oito espécies de Anacampserotaceae e analisou as WBTs do caule e de folhas quanto seu tipo de espessamento e medidas quantitativas de seis espécies. Os resultados apontam a 1) origem estipular das escamas e não protodérmica, como mencionado na literatura; 2) primeiro relato de profilos na família e 3) WBTs ocorrem em Anacampseros e estão ausente em Talinopsis, grupo considerado basal na família. / Anacampserotaceae belongs to the order Caryophyllales, suborder Portulacineae, currently consists of three genus (Anacampseros L., Grahamia Gill ex Hook and Talinopsis A. Gray) formerly circumscribed to the family Portulacaceae. Anacampserotaceae\'s representative presents lignified axillary structures very distinctive, like trichomes, scales and bristles and also WBT cells that is presented only by two other families of Caryophyllales, Aizoaceae e Cactaceae. The axillary characters are very informative for the ACPT clade (Anacampserotaceae, Cactaceae, Portulacaceae and Talinaceae) and has been revealing important data for the phylogenetic studies on the Portulacineae suborder, nevertheless, there is a controversy regarding the Anacampserotaceae axillary structures ontogenetic origin, what may lead to misguided interpretations for the ACPT phylogenetic relationships. Before it, this research have accomplished the ontogenetic study of the structures present on the leaf axil of eight Anacampserotaceae species and examined stem and leaf WBTs, as for its secondary thickening type and quantitative measurements of six species. The results points 1) that at least one structure (scale) has leaf origin instead of protodermic origin, as stated by some authors; 2) first report of prophylls for the family and 3) WBTs occur in Anacampseros but they are absent in Talinopsis, a basal group of Anacampserotaceae.

Anacampserotaceae pertence à ordem Caryophyllales, subordem Portulacineae, atualmente constituída por três gêneros (Anacampseros L., Grahamia Gill ex Hook e Talinopsis A. Gray) antes pertencentes à família Portulacaceae. Os representantes de Anacampserotaceae apresentam estruturas axilares lignificadas muito características, como tricomas, escamas (scales) e cerdas (bristles) e também apresentam células do tipo WBT que ocorre somente em mais outras duas famílias de Caryophyllales, Aizoaceae e Cactaceae. Os caracteres axilares são muito informativos para o clado ACPT (Anacampserotaceae, Cactaceae, Portulacaceae e Talinaceae) e vêm revelando importantes dados aos estudos filogenéticos da subordem Portulacineae, no entanto, há uma controvérsia quanto a origem ontogenética das estruturas axilares de Anacampserotaceae, o que pode levar a interpretações equivocadas das relações filogenéticas do grupo. Diante disso, este trabalho realizou um estudo ontogenético das estruturas presentes na região da axila foliar de oito espécies de Anacampserotaceae e analisou as WBTs do caule e de folhas quanto seu tipo de espessamento e medidas quantitativas de seis espécies. Os resultados apontam a 1) origem estipular das escamas e não protodérmica, como mencionado na literatura; 2) primeiro relato de profilos na família e 3) WBTs ocorrem em Anacampseros e estão ausente em Talinopsis, grupo considerado basal na família. / Anacampserotaceae belongs to the order Caryophyllales, suborder Portulacineae, currently consists of three genus (Anacampseros L., Grahamia Gill ex Hook and Talinopsis A. Gray) formerly circumscribed to the family Portulacaceae. Anacampserotaceae\'s representative presents lignified axillary structures very distinctive, like trichomes, scales and bristles and also WBT cells that is presented only by two other families of Caryophyllales, Aizoaceae e Cactaceae. The axillary characters are very informative for the ACPT clade (Anacampserotaceae, Cactaceae, Portulacaceae and Talinaceae) and has been revealing important data for the phylogenetic studies on the Portulacineae suborder, nevertheless, there is a controversy regarding the Anacampserotaceae axillary structures ontogenetic origin, what may lead to misguided interpretations for the ACPT phylogenetic relationships. Before it, this research have accomplished the ontogenetic study of the structures present on the leaf axil of eight Anacampserotaceae species and examined stem and leaf WBTs, as for its secondary thickening type and quantitative measurements of six species. The results points 1) that at least one structure (scale) has leaf origin instead of protodermic origin, as stated by some authors; 2) first report of prophylls for the family and 3) WBTs occur in Anacampseros but they are absent in Talinopsis, a basal group of Anacampserotaceae.

Wood is a complex organic orthotropic viscoelastic material with acellular structure. When stressed, wood will deform over timethrough a process called creep. Creep affects all wooden structureand can be difficult, time-consuming and expensive to measure. For this thesis, a simple computer model of the woodenmicrostructure was developed. The hypothesis was that the modelledmicrostructure would display similar elastic and viscoelasticproperties as the macroscopic material. The model was designed by finding research with cell geometries ofconiferous trees measured. The model considered late- and earlywoodgeometries as well as growth rings. Rays were ignored as they onlycomposed 5-10% of the material. By applying a finite element method, the heterogeneous late- andearlywood cells could be homogenized by sequentially loading thestrain vector and calculating the average stress. The computer model produced stiff but acceptable values for theelastic properties. Using the standard linear solid method to modelviscoelasticity, the computer model assembled creep curvescomparable to experimental results. With the model sufficiently validated, parametric studies on thecell geometry showed that the elastic and viscoelastic propertieschanged greatly with cell shape. An unconventional RVE was alsotested and shown to give identical result to the standard RVE. Although not perfect, the model can to a certain degree predict theelastic and viscoelastic characteristics for wood given itscellular geometry. Inaccuracies were thought to be caused byassumptions and approximations when building the model.

Most strength grading machines on the European market use an averagemodulus of elasticity (MOE), estimated on a relatively large distance along awood member, as the indicating property (IP) to bending strength. Theaccuracy of such grading machines in terms of coefficient of determination israther low at R2 ≈ 0.5. This research is motivated by a desire to increase theaccuracy of the strength grading in the industry today. The aim of the presentstudy is to contribute knowledge of local variation in bending stiffness/MOEwith high resolution and thus locate weak sections due to stiffness reducingfeatures (the most important is knots) for structural timber.The present study introduces three methods that involve structural dynamics,classical beam theory and optical measurement to assess local wood stiffness.Specifically: The dynamic method, in which a wood member is treated as an ordinaryphysical structure and the local stiffness is studied by exploring itsdynamic properties. In Method II, a bending MOE profile is established based on local fibre angle information. The local fibre orientation is detected through highresolution laser scanning based on the tracheid effect.  For Method III, a bending MOE profile is established using surfacestrain information under four-point bending. A high resolution strainfield is obtained using the digital image correlation (DIC) technique. From the present study, the two latter methods are more favourable inevaluating the local stiffness within a piece of structural timber. Moreover, thestudy reveals that the established bending MOE profiles using the two lattermethods, i.e. based on information of the local fibre angle and surface strain,agree reasonably well. However, for some patterns of knot clusters, the localbending MOE, calculated on the basis of fibre angles, is significantly higherthan the local bending MOE estimated on the basis of surface strain. / De flesta av de utrustningar för hållfasthetssortering som utnyttjas på deneuropeiska marknaden använder ett medelvärde på elasticitetsmodulen(MOE), beräknat på en relativt stor längd av en sågad planka, som indikativparameter (IP). Sådan hållfasthetssortering ger en noggrannhet i termer avförklaringsgrad på R2 ≈ 0.5, vilket är ganska lågt. Arbetet i denna studiemotiveras av en önskan att öka noggrannheten i hållfasthetssorteringen. Syftetmed denna studie är att bidra med kunskap om lokala variationer iböjstyvhet/MOE med hög upplösning och att lokalisera veka snitt (där kvistarär den viktigaste försvagande faktorn) för konstruktionsvirke.Den aktuella studien introducerar tre metoder som omfattar strukturdynamik,klassisk balkteori och optisk mätning vid bedömningen av lokal styvhet imaterialet. Specifikt:  Metod I, där den lokala böjstyvheten studerades genom de dynamiskaegenskaperna såsom egenfrekvens och modform.  Metod II, där en MOE profil beräknas på basis av information om lokalafibervinklar på ett virkesstyckes ytor. Den lokala fiberorienteringen mätsmed högupplöst laserskanning baserad på den så kallade trakeideffekten.  Metod III, där en MOE-profil fastställdes med hjälp avtöjningsinformation för en hel flatsida av en planka belastad med konstantböjmoment. Det högupplösta töjningsfältet erhölls med hjälp av teknikför Digital Image Correlation (DIC). Studien visar att de två sistnämnda metoderna är mycket lämpade för attutvärdera den lokala styvheten i ett virkesstycke. Dessutom visar studien att deMOE-profiler som togs fram med hjälp av de två sistnämnda metoderna,vilka baseras på information om lokala fibervinklar och töjningsfältet på ytan,stämde överens för större delen av virkesstycket. För visa kvistgrupper kan dock den lokala böjstyvheten högre med metoden baserad på fibervinklar.