A Guide to the Biology and Use of Forest Tree Seeds
L A N D M A N A G E M E N T H A N D B O O K
30 A Guide to the Biology and Use of Forest Tree Seeds
1996
Province of British Columbia
SPLENDOR OCCASU Ministry of Forests Research Program SINE A Guide to the Biology and Use of Forest Tree Seeds
Carole Leadem
Province of British Columbia
SPLENDOR OCCASU Ministry of Forests Research Program
SINE Canadian Cataloguing in Publication Data Leadem, Carole Louise Scheuplein, – A guide to the biology of forest tree seeds
(Land management handbook ; )
Includes bibliographical references: p. ---
. Seeds. . Trees - British Columbia - Seeds. . Gymnosperms - British Columbia - Seeds. . Angiosperms - British Columbia - Seeds. . Reforestation . I. British Columbia. Ministry of Forests. Research Branch. II. Title. III. Series.
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Prepared by Carole Leadem B.C. Ministry of Forests Glyn Road Research Station Glyn Road Victoria, BC
for BC Ministry of Forests Research Branch Bastion Square Victoria, BC
Published by B.C. Ministry of Forests Forestry Division Services Branch Production Resources 1205 Broad Street, 2 Floor Victoria, BC
©␣ Province of British Columbia
Copies of this and other Ministry of Forests titles are available from: Crown Publications Inc. Fort Street Victoria, BC ACKNOWLEDGEMENTS
I am grateful to Dr. D. George Edwards, Canadian Izard, Paul Nystedt, and Heather Strongitharm—and Forest Service, for his patience, guidance, and con- Anna Gamble for the publication’s design. structive criticism, and for giving so generously of his The efforts of the editorial team are most greatly wealth of knowledge about tree seeds. appreciated: Dr. Annette Walker, Fran Aitkens, and Thanks are extended to the many B.C. Ministry of Susan Bannerman. Andrew MacKinnon verified the Forests reviewers who shared their expertise and pro- tree species and scientific authorities mentioned in vided useful comments: Rob Bowden-Green, Heather this handbook. Rooke, and Dave Kolotelo of the Tree Seed Centre in I thank the following suppliers of the seed samples Surrey; Karen Yearsley of the Research Branch; Tony used for this publication’s photographs: Don Pigott, Willingdon of the Surrey Nursery; and Clare Hewson Yellow Point Propagation, Ladysmith; Peter Hellenius, of the Interior Seed Orchards in Vernon. Thanks also Silva Enterprises, Prince George; and the Ministry of to Joe Wong of Woodmere Nursey in Telkwa, and Forests Tree Seed Centre, Surrey. Tom Gore of the Candace Laird of the Silviculture Institute of University of Victoria’s Biology Department kindly British Columbia. made available his extensive photographic expertise Joanne Clark provided valuable technical support and facilities. Peggy Frank drew the illustrations for in producing the text and figures, and helped in figures and . Donald Gunn drew the illustrations countless other ways with the final manuscript. for figures and , and the cover topic indicator. I appreciated the work, suggestions, and enthusiasm D. George Edwards supplied the x-ray photos for of the Production Resources staff — especially David Figure .
iii CONTENTS
Acknowledgements ...... iii
Introduction ......
The Basic Principles of Tree Seed Biology ...... . Seed Structure ...... . Development and Maturation ...... . Dormancy ...... . Germination ...... .. Hydration ...... .. Activation of growth processes ...... .. Emergence ...... .. Environmental factors ......
Applying the Principles of Tree Seed Biology ...... . Tree Seed Biology and Reforestation ...... . Seed Quality and Vigour ...... . Seed Collection and Storage ...... . Dormancy ...... . Germination ...... .. Hydration ...... .. Oxygen ...... .. Temperature ...... .. Light ...... .. Other factors ...... . Natural Regeneration ......
Conclusion ...... Appendix . Forest tree species occurring in British Columbia ...... Glossary ...... References ......
v
Dormancy-release treatments for tree seeds ......
Stratification regimes commonly used for conifers grown in British Columbia ......
Moisture content guidelines for tree seeds ......
Forest tree seed anatomy (longitudinal sections) ...... Wings aid in the dispersal of seeds ...... Some trees contain resin vesicles in their seed coats ......
Seeds of the same genus can vary in size and shape ......
Typical development and maturation cycles of British Columbia conifer seeds ...... Comparison of the major steps in the natural and artificial regeneration sequences of forest tree seedlings ...... Stages of germinant development ...... Absorption of far-red light converts the pigment phytochromefar-red back to phytochromered ...... Vigorous seeds complete germination first ......
Mature and immature embryos of Douglas-fir ...... The longevity of seeds increases as seed moisture content and storage temperature decreases ...... Effects of stratification regime on the germination of western hemlock seeds ...... Effects of stratification regime on the germination rates of Pacific silver fir seeds ......
Respiration of subalpine fir seeds during stratification ...... Germination of a) lodgepole pine, b) Sitka spruce, and c) Douglas-fir seeds at different temperatures after stratification for , , , and weeks ...... X-rays are used to determine whether seeds are fully developed, damaged, or have been attacked by insects ......
A young whitebark pine seedling struggles to establish in a high alpine meadow ......
Photographic tableau of forest tree seeds ......
vi 1 INTRODUCTION
The reasons for an interest in forest tree seeds vary widely. Nursery workers, silviculturists, seed orchard managers, cone collectors, and seed dealers have a very practical need for knowledge. But many others have developed a general interest in seed biology because they want to achieve a better under- standing of the natural world around them. Seed maturation, dormancy, and germination are still not completely understood. It remains somewhat of a mystery how a seed can remain viable for many years in the forest duff, then, responding to some cue, break through its woody seed coat and establish itself as an independent seedling. However, we know some of the factors critical to those processes, and we know that the effects of these factors may vary, depending on the physiological state of the seed. At the moment of natural seedfall, the potential quality of seeds is as high as it will ever be. To main- tain that quality and to produce the best seedlings for reforestation, knowledge of tree seed biology is essential. This handbook describes the basic principles that govern the biology of forest tree seeds and examines how these principles might apply to reforestation. Its intent is to give an overall picture of how and why seeds may germinate and to provide some understanding of a remarkable process.
2 THE BASIC PRINCIPLES OF TREE SEED BIOLOGY
. Seed Structure
A seed is a unique package containing the essential provides the energy supplies and structures of a new seedling and the nutrients to raw materials needed by the germinating embryo. support early growth. This package is constructed This tissue maintains the developing seedling until during a maturation period, after which the seed its photosynthetic and water uptake systems are undergoes a period of dormancy, followed by a able to support it. It contains vitamins, plant growth reactivation process referred to as germination. regulators, minerals, and many organic compounds, Each step in the sequence is critical to optimum all essential for normal embryo growth. seed performance. Nutritive tissues of conifer (gymnosperm) tree A fully developed seed consists of an embryo sur- seeds, and broad-leaved (angiosperm) tree seeds rounded by nutritive tissue, all of which is enclosed differ in several important respects, although both in a protective seed coat. The anatomy of several types of tissue perform the same function. tree seeds is shown in Figure . In conifers, the nutritive tissue is referred to as is a plant in miniature, containing the megagametophyte, and contains a large pro- rudimentary versions of the basic structures needed portion of fats and proteins. The tissue derives by the new seedling for growth and development: entirely from the female parent and has a single primary leaves (cotyledons), primary root (radicle), chromosome complement (). The nutritive the stem below the cotyledons (hypocotyl), and the tissue of conifer seeds is physically separate, and stem above the cotyledons (epicotyl). food must move from the megagametophyte to the Growth regions (meristems) located at the base embryo by diffusion. of the cotyledons and behind the root tip are the Nutritive tissue of broad-leaved tree seeds is source of new cells for the shoot and root growth called the endosperm and carries a chromosome of the seedling. complement of . It is produced from the union of The embryo contains the genetic makeup of the one set of chromosomes derived from the male par- new seedling. The embryo is the product of fertiliz- ent, and of two sets derived from the female parent. ation, or the union of the egg from the female par- Many broad-leaved tree seeds store the major part ent with the sperm contained in the male pollen. of their food supply in the cotyledons. The structure Both the female parent and male parent contribute of angiosperm seeds allows for the direct transfer of a single chromosome complement to the egg and food supplies from the endosperm or cotyledons be- the sperm. A mature seed, therefore, has a , or cause they are physically attached to the embryo. double chromosome complement. 2.7 mm 9 mm Seed wing
Pericarp Seed coat
Seed coat Cotyledons
Cotyledon Hypocotyl Embryo Embryo Hypocotyl Megagametophyte
Radicle Radicle
Micropyle Micropyle
Forest tree seed anatomy (longitudinal sections): red alder, an angiosperm (left); and Douglas-fir, a gymnosperm (right).
Words in bold are defined in the glossary.
a) Seed length: 2.6 cm b) Seed length: 2.1 cm c) Seed length: 1.0 cm d) Seed width: 0.5 cm e) Total seed length (seed = 1 cm; wing = 1.6 cm) (seed = 0.7 cm; wing = 1.3 cm) (seed = 0.3 cm; wing = 0.7 cm) (seed = 5 mm) (with wings): 5.2 cm Wings aid in the dispersal of seeds: a)␣ Pacific silver fir; b)␣ ponderosa pine; c) white spruce; d) yellow-cedar; e) bigleaf maple. Average dimensions of seeds provided.
of the seed (testa) provides area in the coat. During germination, embryo elon- physical protection for the embryo and nutritive gation and degradation of nutritive tissue assist the tissue. As well, it regulates the movement of water, radicle to emerge through the micropyle. Seeds may oxygen, and carbon dioxide in and out of the seed. vary greatly in size, colour, and shape. Even seeds In many species, the seed coat has membranous of closely related species can appear very different protrusions called “wings” that enhance the wind (Figure 4). dispersal of mature seeds (Figure ). In species such as pine,2 hemlock, and spruce, the wings may be . Development and Maturation easily detached, but in species such as western redcedar and yellow-cedar, the wings cannot be During maturation, a fertilized ovule is transformed removed without damaging the seed. Conifers such into a fully developed seed, containing all the ele- as western redcedar, the true firs, western hemlock, ments to produce a new tree. In the early stages of and mountain hemlock contain resin vesicles in the reproductive cycle, ovules develop in female their seed coats (Figure ). These can be damaged by cones and pollen develops in male cones. Pollen is improper handling, resulting in reduced viability of released in the spring and carried by the wind from the seed. the male to the female cones. In most British The seed coat has a chromosome complement of Columbia conifers, fertilization takes place during 2. It develops from the tissues (integuments) of the spring or early summer, shortly after pollination. the female parent tree that surround the ovule be- Following fertilization, the embryo grows until, at fore fertilization. The opening (micropyle) through maturity, it occupies almost the full length of the which pollen enters the ovule remains as a weak seed.
v v
v
v v
v a) b) c)
Some trees contain resin vesicles in their seed coats: a)␣ western redcedar, seed length = 0.5 cm; b)␣ Pacific silver fir, seed length = 1.1 cm; c)␣ western hemlock, seed length = 0.4 cm.
Scientific and common names of forest tree species occurring in British Columbia are listed in Appendix 1.
For Douglas-fir, redcedar, spruces, true firs, and hemlocks, the development and maturation cycle takes about months (Figure ). In pines, complete development takes months because fertilization is delayed for one year after pollination. In yellow- cedar, pollination and fertilization take place in the same growing season, but the total cycle usually lasts about months. Dehydration is an essential part of the maturation process. Water is lost from the seed, cell membranes assume a more condensed form, and physiological processes such as respiration diminish to very low levels. Simple compounds are changed to starches, fats, and proteins. These complex compounds can remain stable over many years, enabling seeds to lie dormant or to be stored for long periods. During germination the process is reversed as Seeds of the same genus can vary in size and storage compounds are broken down into simpler shape (counterclockwise from top): Korean pine, Coulter pine, limber pine, whitebark pine, forms (such as sugars and amino acids) that can be ponderosa pine, western white pine, Caribbean easily used by the embryo. Mature seeds are released pine, jack pine (2 seeds), lodgepole pine. naturally from cones in late summer and fall. Depending on the species, dispersal sometimes would leave vulnerable seedlings exposed to harsh continues into the following spring. winter conditions. In nature, dormant seeds remain inactive until favourable growing conditions occur . Dormancy the following spring. Some may remain dormant for two growing seasons or more. Seeds can maintain In the seeds of many tree species, maturation is ac- viability for many years in a dormant state. companied by the induction of a state of dormancy. Seeds are released from dormancy through This is an advantage for seeds that mature in late changes that occur during their exposure to cold, wet summer to early fall, since immediate germination conditions over winter, and they (usually) germinate
Reproductive cycles Year 1 Year 2 Year 3
Bud initiation Mature seeds
Douglas-fir, redcedar spruces, true firs, and hemlocks