Chapter 1
Introduction
PURPOSE
Each chapter in this book begins with a brief summary of
its purpose(s); thus, in keeping with this format, one
objective of Chapter 1 is to explain the philosophy for the
whole book. The original concept for this book was to
synthesize available information related to seed germina-
tion ecology, including evolutionary aspects. However, as
the mass of information we were collecting on seed dor-
mancy and germination continued to increase, it became
clear that we should use this database to (1) determine if
plants with specialized life cycle and/or habitats were
unique with regard to seed dormancy and germination
characteristics, (2) obtain an understanding of the world
biogeography of whole-seed dormancy and (3) formulate
hypotheses concerning the origin and evolutionary rela-
tionships of the various kinds of dormancy in seeds.
Thus, this book attempts to cover the present state of
knowledge of the ecology, biogeography and evolution
of seed dormancy and germination.
Other objectives of this introductory chapter are to
explain what we mean by seed germination ecology,
provide an overview of the general kinds of information
needed to understand the seed germination ecology of a
species and comment on the values of such studies. Also,
attention will be given to reasons why this book on seed
germination is needed, and how it differs from previous
ones written on the subject.
SEED GERMINATION ECOLOGY
Much information is available in the literature on the seed
germination phenology of individual species, and these data
show that many species have a characteristic germination
season (or seasons). For many species, the time of year
when germination is possible is quite limited, e.g., only
in autumn, spring or the wet season. In contrast, the germi-
nation season for other species is long, e.g., throughout the
growing season or even throughout the whole year. Thus,
the broad objective of a seed germination ecologist is to
explain how the timing of germination is controlled in
nature, and the ecological and evolutionary origins and
consequences of this timing.
C.C. Baskin and J.M. Baskin: Seeds, Second Edition. DOI: http://dx.doi.org/10.1016/B978-0-12-416677-6.00001-9
© 2014 Elsevier Inc. All rights reserved.
From a careful reading of Evenari’s (1980/81) paper
on the history of germination research, one comes to the
conclusion that Theophrastus (c. 372 287 B.C.) should be
called the Father of Seed Germination Ecology. This
remarkable early Greek philosopher and scientist knew that:
(1) food reserves are stored in seeds, (2) the environmental
conditions under which seeds mature affect their germina-
tion characteristics and (3) germination can be influenced
by climatic factors, inhibitors, seed age and seed coats.
Determining what controls the timing of seed germi-
nation in the field requires information on the seed,
environmental conditions in the habitat and how the two
interact from time of seed maturation to germination. The
best way to study the seed germination ecology of a
species is to break the problem into a series of questions:
(1) When do seeds mature? (2) When are they dispersed?
(3) What is the natural germination unit, i.e., seed, fruit
or fruit with accessory parts? (4) What is the dormancy
state of seeds at the time of maturation and at dispersal?
(5) What are the environmental conditions in the habitat
between time of maturation and germination? (6) What
environmental conditions are required to break dormancy
and to induce it? (7) What conditions are required to
promote germination of nondormant seeds?
In the study of seed germination ecology, other kinds
of questions come to mind. For example, how do genetics
and/or the climate under which seeds develop influence
the requirements for dormancy break and germination?
How much variation is there within a species? Does the
species form a long-lived seed bank? Do buried seeds
undergo changes in their dormancy state? What role does
timing of germination play in a plant’s fitness?
Since the days of Theophrastus, a vast amount of
information related to seed germination has accumulated;
however, as can be seen in subsequent chapters, much
remains to be learned about some aspects of the field.
There are two primary reasons why people are interested
in learning how the timing of seed germination is con-
trolled under natural conditions: economics and aca-
demics. These two reasons for doing seed research are
not independent, and discoveries by people with one point
of view frequently benefit those approaching the subject
from the other way.
1
2 Seeds
Information on seed germination potentially has great
monetary value. A knowledge of what controls the timing
of germination enhances (1) planning for effective control
of weeds, (2) propagation of native economically impor-
tant trees, shrubs, vines, forbs and grasses and (3) recla-
mation of damaged ecosystems. Thus, much has been
learned about seed germination from the research efforts
of agronomists, foresters, rangeland specialists, reclama-
tion ecologists and weed scientists.
Information on seed germination also is important
because it contributes to a better understanding of certain
biological concepts such as plant reproductive strategies,
life history traits, adaptation to habitats and physiological
and molecular processes. Considerable research attention
has been given to seed development and maturation, storage
and mobilization of food reserves, seed storage behavior—
especially of economically important plants, aging, cellular,
biochemical molecular events in germination and responses
of nondormant seeds to various environmental factors, such
as temperature, light, soil moisture, nutrients and salinity.
Thus, much has been (and continues to be) learned about
seed germination from the research of plant anatomists, bio-
chemists, ecologists, geneticists, physiologists and molecular
biologists.
A BOOK ON SEED DORMANCY AND
GERMINATION FROM A DIFFERENT
PERSPECTIVE
Need
Since information on seed germination is derived from
the research efforts of people in a wide variety of disci-
plines, it should not be surprising that the data are pub-
lished in many different kinds of journals, in countries
all over the world and in a variety of languages. Thus,
the general subject matter of journals that publish papers
dealing with some aspect of seed germination ranges
from weed science, agronomy, crop science, range
management, horticulture, forestry, botany, ecology,
conservation biology, morphology, genetics, physiology,
biochemistry and molecular biology. However, a few
journals, e.g., Journal of Seed Technology, Seed Science
and Technology, Seed Science Research, Journal of New
Seeds, Journal of Seed Research and Revista Brasileira
de Sementes, are devoted exclusively to seed research.
Information on seed germination also is found in agricul-
ture research station bulletins, dissertations and theses,
proceedings of symposia and conferences (e.g., Nicolas
et al., 2003; Adkins et al., 2007), as well as in reviews
(e.g., Annual Review of Ecology Evolution and Systematics,
Annual Review of Plant Physiology and Encyclopedia of
Plant Physiology) and in books.
Some books on seeds emphasize (1) chemical composi-
tion, physiology and biochemistry (Khan, 1977, 1982;
Bewley and Black, 1982, 1983, 1994; Labouriau, 1983;
Murray, 1984; Zhang, 2004; Black et al., 2006; Bewley
et al., 2012), (2) types of dormancy, and how they are broken
(Nikolaeva, 1969; Bradbeer, 1988), (3) structure (Corner,
1976), (4) morphology (Davis, 1993) and (5) seed technol-
ogy (Copeland and McDonald, 2001; Basra, 2006). Others
cover various aspects of seed physiology and/or dormancy
and how environmental factors affect germination (Crocker
and Barton, 1957; Heydecker, 1972; Kozlowski, 1972;
Fenner, 1985; Mayer and Poljakoff-Mayber, 1989; Kigel and
Galili, 1995; Benech-Arnold and Sanchez, 2004). Books
have been written (or compiled) on special topics such as
embryo and seed development and germination
(Obroucheva, 1999; Batygina, 2006); seed aging (Priestley,
1986); identification (Bouman et al., 2000; Lentz and
Dickau, 2005; Sweedman and Merritt, 2006); conservation
(Smith et al., 2003; Guerrant et al., 2004); ecology (Fenner
and Thompson, 2005); storage and longevity (Roberts,
1972a; Sacande et al., 2004); testing (Elias et al., 2012); soil
seed banks (Leck et al., 1989; Thompson et al. 1997); germi-
nation of desert plants (Gutterman, 1993, 2002), flower seeds
(McDonald and Kwong, 2005) and grasses (Simpson, 1990);
restoration (Munoz et al., 1999; Elliott et al., 2005;
L’Huillier et al., 2010); and role of the seeds in regeneration
of plant communities (Fenner, 1992a, 2000). It is not uncom-
mon for books to contain one or more chapters on the germi-
nation of special groups of plants such as orchids (Arditti,
1977a, 1982; Rasmussen, 1995), grasses (Cheplick, 1998),
halophytes (Ungar, 1991b), parasites (Parker and Riches,
1993), trees (Farmer, 1997), weeds (Benech-Arnold and
Sanchez, 1995) or plants subjected to flooding (Scarano
and Franco, 1998). A number of handbooks have been pub-
lished that provide detailed instructions on how to germinate
seeds of weeds (Andersen, 1968), woody plants (Young and
Young, 1992; Compilation Committee, 2000; Schmidt,
2000; Vozzo, 2002; Piotto and Di Noi, 2003; Bonner and
Karrfalt, 2008), native plants (e.g., Langkamp, 1987;
Culliney and Koebele, 1999; Cullina, 2000, 2002b; Lilleeng-
Rosenberger, 2005) and plants of interest to gardeners
(Deno, 1994).
Although much information obviously was available
on seed germination ecology, it was scattered throughout
the scientific literature. Thus, since no single comprehen-
sive source of data was available on the subject, we under-
took the task of doing a synthesis and published the first
edition of our seed book in 1998, which was issued in
paperback in 2001. As we continued to learn about seeds
after July 1998, it quickly became clear that our “synthe-
sis” had not been totally successful—i.e., there was still a
lot more information that needed to be added to the book.
Thus, we have tried to meet the challenge by adding large
quantities of new information to Seeds.
Chapter | 1 Introduction
A comprehensive synthesis of seed ecology will:
(1) facilitate and help focus research efforts, (2) identify
questions that need to be addressed by additional research,
(3) make it possible to analyze the presence of the various
kinds of dormancy in different life forms of plants (i.e.,
trees, shrubs, lianas and herbaceous species) in the major
vegetation types of the world and (4) provide a foundation
to which other kinds of information (such as fossil history
and phylogenetic positions of families) can be added to
formulate hypotheses on the evolution of the various kinds
of seed dormancy.
Content
Our book differs from others on seed germination in
seven important ways: (1) It contains a chapter on
procedures for doing germination studies so that labora-
tory and greenhouse results can be extrapolated to the
field; (2) The use of a hierarchical system of classification
of seed dormancy is discussed, along with the benefit of
using Nikolaeva’s formula system for dormancy classifi-
cation as a means of facilitating international communica-
tion about the kind of seed dormancy being investigated;
(3) Information on many topics—e.g., dormancy-breaking
requirements of seeds with each class of dormancy and
genetics of seed dormancy and germination—is placed
into an ecological context; (4) A critical evaluation of
methodology used in soil seed bank studies of plant com-
munities is presented, and it shows that many researchers
have sampled mixtures of persistent and transient seed
banks rather than persistent seed banks only; (5) Data
were compiled on species with different life forms grow-
ing in the major vegetation types worldwide to gain a
global perspective on geographical/ecological relationships
of the dormancy-breaking and germination requirements of
seeds; (6) Data on environmental conditions required to
break seed dormancy and stimulate germination have been
3
synthesized for species with specialized life cycles and/or
habitats, including parasites, saprophytes, orchids, carnivo-
rous plants, aquatics, halophytes and psammophytes, and
this information has been put into an ecological context;
(7) For the first time, an attempt has been made to unravel
the evolutionary/phylogenetic origins and relationships of
the various kinds of seed dormancy and to explain the
conditions under which each may have evolved.
The revision of our book has led to the expansion of
many topics such as the definition of seed dormancy,
abscisic acid (ABA) and development of seed dormancy,
phytochromes, hydrothermal time models, inbreeding
depression and germination, animal predation of seeds,
the two-step requirement for breaking physical dormancy,
water gaps in seeds with physical dormancy and how they
function and germination ecology of tropical montane
species. The database for the world biogeography of
seed dormancy has been expanded from 3580 to about
13,600 species, and the Takhtajan phylogenetic scheme
for angiosperms has been replaced by APG III (Chase
and Reveal, 2009). Finally, each chapter now contains
one to many new topics such as need to check seeds for
presence of embryo; new plant comes from an undifferen-
tiated embryo that develops into a differentiated embryo;
dichotomous key for dormancy classes; physiological epi-
cotyl dormancy; orchid seed food reserves and asymbiotic
germination; nondeep simple epicotyl morphophysiologi-
cal dormancy; sensitivity cycling in seeds with physical
dormancy; spatial distribution of seeds in the soil seed
bank and role of seed banks in restoration of plant com-
munities; testing dormancy evolution hypotheses; seed
germination in relation to parental effects, pollen compe-
tition, local adaptation, climate change, partial predation
of the seeds (and they still germinate), biological crusts,
nurse plants and karrikinolide in smoke from burning
plants; and seed germination of rare and of invasive
species.