D.J. Durzan – författare

For many, the terms aging, maturation and senescence are synonymous and used interchangeably, but they should not be. Whereas senescence represents an endogenously controlled degenerative programme leading to plant or organ death, genetiC aging encompasses a wide array of passive degenerative genetiC processes driven primarily by exogenous factors (Leopold, 1975). Aging is therefore considered a consequence of genetiC lesions that accumulate over time, but by themselves do not necessarily cause death. These lesions are probably made more severe by the increase in size and complexity in trees and their attendant physiology. Thus while the withering of flower petals following pollination can be considered senescence, the loss of viability of stored seeds more clearly represents aging (Norden, 1988). The very recent book "Senescence and Aging in Plants" does not discuss trees, the most dominant group of plants on the earth. Yet both angiospermic and gymnospermic trees also undergo the above phenomena but less is known about them. Do woody plants senesce or do they just age? What is phase change? Is this synonymous with maturation? While it is now becoming recognized that there is no programmed senescence in trees, senescence of their parts, even in gymnosperms (e. g. , needles of temperate conifers las t an average of 3. 5 years), is common; but aging is a readily acknowledged phenomenon. In theory, at least, in the absence of any programmed senescence trees should -live forever, but in practice they do not.

For many, the terms aging, maturation and senescence are synonymous and used interchangeably, but they should not be. Whereas senescence represents an endogenously controlled degenerative programme leading to plant or organ death, genetiC aging encompasses a wide array of passive degenerative genetiC processes driven primarily by exogenous factors (Leopold, 1975). Aging is therefore considered a consequence of genetiC lesions that accumulate over time, but by themselves do not necessarily cause death. These lesions are probably made more severe by the increase in size and complexity in trees and their attendant physiology. Thus while the withering of flower petals following pollination can be considered senescence, the loss of viability of stored seeds more clearly represents aging (Norden, 1988). The very recent book "Senescence and Aging in Plants" does not discuss trees, the most dominant group of plants on the earth. Yet both angiospermic and gymnospermic trees also undergo the above phenomena but less is known about them. Do woody plants senesce or do they just age? What is phase change? Is this synonymous with maturation? While it is now becoming recognized that there is no programmed senescence in trees, senescence of their parts, even in gymnosperms (e. g. , needles of temperate conifers las t an average of 3. 5 years), is common; but aging is a readily acknowledged phenomenon. In theory, at least, in the absence of any programmed senescence trees should -live forever, but in practice they do not.

2. IMPORTANCE OF NITROGEN METABOLISM 2. 1. Range of naturally occurring nitrogenous components in forest trees 2. 2. Gene expression and mapping 2. 3. Metabolic changes in organized and unorganized systems 2. 4. Nitrogen and nutrition 2. 5. Aspects of intermediary nitrogen metabolism 3. NITROGEN METABOLISM IN GROWTH AND DEVELOPMENT 3. 1. Precultural factors 3. 2. Callus formation 3. 3. Cell suspensions 3. 3. 1. Conifers 3. 3. 2. Acer 3. 4. Morphogenesis 3. 4. 1. Nitrogen metabolism of natural embryos 3. 4. 2. Somatic embryogenesis 3. 4. 2. 1. Sweetgum (Liquidambar styraciflua) 3. 4. 2. 2. Douglar-fir and loblolly pine 3. 4. 3. Organogenesis 4. OUTLOOK 11. CARBOHYDRATE UTILIZATION AND METABOLISM - T. A. Thorpe 325 1. INTRODUCTION 2. NUTRITIONAL ASPECTS 3. CARBOHYDRATE UPTAKE 4. CARBOHYDRATE METABOLISM 4. 1. Sucrose degradation 4. 2. Metabolism of other carbon sources 4. 3. Hexose mobilization and metabolism 4. 3. 1. Cell cycle studies 4. 3. 2. Growth studies 4. 3. 3. Organized development 4. 4. Cell wall biogenesis 4. 4. 1. Primary cell walls 4. 4. 2. Cell wall turnover 4. 4. 3. Secondary cell walls 4. 5. Carbon skeleton utilization 5. OSMOTIC ROLE 6. CONCLUDING THOUGHTS 369 12. THE USE OF IN VITRO TECHNIQUES FOR GENETIC MODIFICATIO~FOREST TREES - E. G. Kirby 1. INTRODUCTION 2. IN VITRO SELECTION 2. 1. Natural variation 2. 2. Induction of variation 2. 3. Selection techniques 2. 4. Plant regeneration 2 . • 5. Applications x 3. SOMATIC HYBRIDIZATION 3. 1.

Since the first edition of our book "Tissue Culture in Fores­ try" in 1982 we have witnessed remarkable advances in cell and tissue culture technologies with woody perennials. In addition to forest biologists in government, industry, and universities, we now have molecular biologists, genetic engineers, and biochemists using cell and tissue cultures of woody species routinely. There­ fore, the time has come for an update of the earlier edition. In our present effort to cover new developments we have expanded to three volumes: 1. General principles and Biotechnology 2. Specific Principles and Methods: Growth and Development 3. Case Histories: Gymnosperms, Angiosperms and Palms The scientific barriers to progress in tree improvement are not so much lack of foreign gene expression in plants but our current inabili ty to regenerate plants in true-to-type fashion on a mas­ sive and economic scale. To achieve this in the form of an appro­ pr iate biotechnology, cell and tissue culture will increasing ly require a better understanding of basic principles in chemistry and physics that determine structural and functional relationships among molecules and macromolecules (proteins, RNA, DNA) within cells and tissues. These principles and their relationship with the culture medium and its physical environment, principles of clonal propagation, and genetic variation and ultrastructure are discussed in volume one.

2. 2. Plant materials 2. 3. Pregrowth conditions 2. 4. Cryoprotectant treatment 2. 5. Freezing 2. 5. 1. Slow freezing 2. 5. 2. Rapid freezing 2. 5. 3. Droplet freezing 2. 6. Storage 2. 7. Thawing 2. 8. Viability testing 2. 9. Post-thaw regrowth 3. EXAMPLES OF CRYOPRESERVATION OF WOODY PLANT MATERIAL 4. POTENTIAL APPLICATION OF CRYOPRESERVATION IN TREE IMPROVEMENT 17. NURSERY HANDLING OF PROPAGULES - J. A. Driver, and 320 G. R. L. Suttle 1. INTRODUCTION 2. COMMERCIAL NURSERY NEEDS VS. LABORATORY PRACTICE 3. SEASONALITY OF GROWTH AND PRODUCTION CYCLES 4. MICROPROPAGATION OPTIONS 4. 1. Trends in commercial micropropagation 4. 1. 1. Contract micropropagation 5. FACTORS AFFECTING SURVIVAL AND GROWTH 5. 1. Hardening of propagules in vitro 5. 2. Greenhouse considerationS------ 5. 3. Field planting 5. 4. New approaches: Direct field rooting 5. 4. 1. Pretreatment in vitro 5. 4. 2. Root induction 5. 4. 3. Field placement 18. MYCORRHIZAE - R. K. Dixon, and D. H. Marx 336 1. INTRODUCTION 2. ROLE OF MYCORRHIZAE IN TREE GROWTH AND DEVELOPMENT 3. PRODUCTION AND APPLICATION OF ECTOMYCORRHIZAL FUNGUS INOCULUM 3. 1. Bareroot stock 3. 2. Container-grown stock 4. FIELD TRIALS WITH ECTOMYCORRHIZAL PLANTING STOCK 5. PRODUCTION AND APPLICATION OF ENDOMYCORRHIZAL INOCULUM 6. FIELD TRIALS WITH ENDOMYCORRHIZAL 7. RESEARCH OPPORTUNITIES 8. SUMMARY 351 19. TISSUE CULTURE APPLICATIUN TO FOREST PATHOLOGY AND PEST CONTROL - A. M. Diner, and D. F. Karnosky 1. INTRODUCTION 2. HOST AND PATHOGEN: CULTURE AND CHALLENGE 2. 1.

2. IMPORTANCE OF NITROGEN METABOLISM 2. 1. Range of naturally occurring nitrogenous components in forest trees 2. 2. Gene expression and mapping 2. 3. Metabolic changes in organized and unorganized systems 2. 4. Nitrogen and nutrition 2. 5. Aspects of intermediary nitrogen metabolism 3. NITROGEN METABOLISM IN GROWTH AND DEVELOPMENT 3. 1. Precultural factors 3. 2. Callus formation 3. 3. Cell suspensions 3. 3. 1. Conifers 3. 3. 2. Acer 3. 4. Morphogenesis 3. 4. 1. Nitrogen metabolism of natural embryos 3. 4. 2. Somatic embryogenesis 3. 4. 2. 1. Sweetgum (Liquidambar styraciflua) 3. 4. 2. 2. Douglar-fir and loblolly pine 3. 4. 3. Organogenesis 4. OUTLOOK 11. CARBOHYDRATE UTILIZATION AND METABOLISM - T. A. Thorpe 325 1. INTRODUCTION 2. NUTRITIONAL ASPECTS 3. CARBOHYDRATE UPTAKE 4. CARBOHYDRATE METABOLISM 4. 1. Sucrose degradation 4. 2. Metabolism of other carbon sources 4. 3. Hexose mobilization and metabolism 4. 3. 1. Cell cycle studies 4. 3. 2. Growth studies 4. 3. 3. Organized development 4. 4. Cell wall biogenesis 4. 4. 1. Primary cell walls 4. 4. 2. Cell wall turnover 4. 4. 3. Secondary cell walls 4. 5. Carbon skeleton utilization 5. OSMOTIC ROLE 6. CONCLUDING THOUGHTS 369 12. THE USE OF IN VITRO TECHNIQUES FOR GENETIC MODIFICATIO~FOREST TREES - E. G. Kirby 1. INTRODUCTION 2. IN VITRO SELECTION 2. 1. Natural variation 2. 2. Induction of variation 2. 3. Selection techniques 2. 4. Plant regeneration 2 . • 5. Applications x 3. SOMATIC HYBRIDIZATION 3. 1.

Since the first edition of our book "Tissue Culture in Fores­ try" in 1982 we have witnessed remarkable advances in cell and tissue culture technologies with woody perennials. In addition to forest biologists in government, industry, and universities, we now have molecular biologists, genetic engineers, and biochemists using cell and tissue cultures of woody species routinely. There­ fore, the time has come for an update of the earlier edition. In our present effort to cover new developments we have expanded to three volumes: 1. General principles and Biotechnology 2. Specific Principles and Methods: Growth and Development 3. Case Histories: Gymnosperms, Angiosperms and Palms The scientific barriers to progress in tree improvement are not so much lack of foreign gene expression in plants but our current inabili ty to regenerate plants in true-to-type fashion on a mas­ sive and economic scale. To achieve this in the form of an appro­ pr iate biotechnology, cell and tissue culture will increasing ly require a better understanding of basic principles in chemistry and physics that determine structural and functional relationships among molecules and macromolecules (proteins, RNA, DNA) within cells and tissues. These principles and their relationship with the culture medium and its physical environment, principles of clonal propagation, and genetic variation and ultrastructure are discussed in volume one.

2. 2. Plant materials 2. 3. Pregrowth conditions 2. 4. Cryoprotectant treatment 2. 5. Freezing 2. 5. 1. Slow freezing 2. 5. 2. Rapid freezing 2. 5. 3. Droplet freezing 2. 6. Storage 2. 7. Thawing 2. 8. Viability testing 2. 9. Post-thaw regrowth 3. EXAMPLES OF CRYOPRESERVATION OF WOODY PLANT MATERIAL 4. POTENTIAL APPLICATION OF CRYOPRESERVATION IN TREE IMPROVEMENT 17. NURSERY HANDLING OF PROPAGULES - J. A. Driver, and 320 G. R. L. Suttle 1. INTRODUCTION 2. COMMERCIAL NURSERY NEEDS VS. LABORATORY PRACTICE 3. SEASONALITY OF GROWTH AND PRODUCTION CYCLES 4. MICROPROPAGATION OPTIONS 4. 1. Trends in commercial micropropagation 4. 1. 1. Contract micropropagation 5. FACTORS AFFECTING SURVIVAL AND GROWTH 5. 1. Hardening of propagules in vitro 5. 2. Greenhouse considerationS------ 5. 3. Field planting 5. 4. New approaches: Direct field rooting 5. 4. 1. Pretreatment in vitro 5. 4. 2. Root induction 5. 4. 3. Field placement 18. MYCORRHIZAE - R. K. Dixon, and D. H. Marx 336 1. INTRODUCTION 2. ROLE OF MYCORRHIZAE IN TREE GROWTH AND DEVELOPMENT 3. PRODUCTION AND APPLICATION OF ECTOMYCORRHIZAL FUNGUS INOCULUM 3. 1. Bareroot stock 3. 2. Container-grown stock 4. FIELD TRIALS WITH ECTOMYCORRHIZAL PLANTING STOCK 5. PRODUCTION AND APPLICATION OF ENDOMYCORRHIZAL INOCULUM 6. FIELD TRIALS WITH ENDOMYCORRHIZAL 7. RESEARCH OPPORTUNITIES 8. SUMMARY 351 19. TISSUE CULTURE APPLICATIUN TO FOREST PATHOLOGY AND PEST CONTROL - A. M. Diner, and D. F. Karnosky 1. INTRODUCTION 2. HOST AND PATHOGEN: CULTURE AND CHALLENGE 2. 1.

2. 2. Plant materials 2. 3. Pregrowth conditions 2. 4. Cryoprotectant treatment 2. 5. Freezing 2. 5. 1. Slow freezing 2. 5. 2. Rapid freezing 2. 5. 3. Droplet freezing 2. 6. Storage 2. 7. Thawing 2. 8. Viability testing 2. 9. Post-thaw regrowth 3. EXAMPLES OF CRYOPRESERVATION OF WOODY PLANT MATERIAL 4. POTENTIAL APPLICATION OF CRYOPRESERVATION IN TREE IMPROVEMENT 17. NURSERY HANDLING OF PROPAGULES - J. A. Driver, and 320 G. R. L. Suttle 1. INTRODUCTION 2. COMMERCIAL NURSERY NEEDS VS. LABORATORY PRACTICE 3. SEASONALITY OF GROWTH AND PRODUCTION CYCLES 4. MICROPROPAGATION OPTIONS 4. 1. Trends in commercial micropropagation 4. 1. 1. Contract micropropagation 5. FACTORS AFFECTING SURVIVAL AND GROWTH 5. 1. Hardening of propagules in vitro 5. 2. Greenhouse considerationS------ 5. 3. Field planting 5. 4. New approaches: Direct field rooting 5. 4. 1. Pretreatment in vitro 5. 4. 2. Root induction 5. 4. 3. Field placement 18. MYCORRHIZAE - R. K. Dixon, and D. H. Marx 336 1. INTRODUCTION 2. ROLE OF MYCORRHIZAE IN TREE GROWTH AND DEVELOPMENT 3. PRODUCTION AND APPLICATION OF ECTOMYCORRHIZAL FUNGUS INOCULUM 3. 1. Bareroot stock 3. 2. Container-grown stock 4. FIELD TRIALS WITH ECTOMYCORRHIZAL PLANTING STOCK 5. PRODUCTION AND APPLICATION OF ENDOMYCORRHIZAL INOCULUM 6. FIELD TRIALS WITH ENDOMYCORRHIZAL 7. RESEARCH OPPORTUNITIES 8. SUMMARY 351 19. TISSUE CULTURE APPLICATIUN TO FOREST PATHOLOGY AND PEST CONTROL - A. M. Diner, and D. F. Karnosky 1. INTRODUCTION 2. HOST AND PATHOGEN: CULTURE AND CHALLENGE 2. 1.

Since the first edition of our book "Tissue Culture in Fores­ try" in 1982 we have witnessed remarkable advances in cell and tissue culture technologies with woody perennials. In addition to forest biologists in government, industry, and universities, we now have molecular biologists, genetic engineers, and biochemists using cell and tissue cultures of woody species routinely. There­ fore, the time has come for an update of the earlier edition. In our present effort to cover new developments we have expanded to three volumes: 1. General principles and Biotechnology 2. Specific Principles and Methods: Growth and Development 3. Case Histories: Gymnosperms, Angiosperms and Palms The scientific barriers to progress in tree improvement are not so much lack of foreign gene expression in plants but our current inabili ty to regenerate plants in true-to-type fashion on a mas­ sive and economic scale. To achieve this in the form of an appro­ pr iate biotechnology, cell and tissue culture will increasing ly require a better understanding of basic principles in chemistry and physics that determine structural and functional relationships among molecules and macromolecules (proteins, RNA, DNA) within cells and tissues. These principles and their relationship with the culture medium and its physical environment, principles of clonal propagation, and genetic variation and ultrastructure are discussed in volume one.