See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/272790378

Seed germination and sowing options

Chapter · January 2014

CITATION READS
1 1,233

3 authors, including:

Tara Luna R. Kasten Dumroese

33 PUBLICATIONS 120 CITATIONS

US Forest Service
273 PUBLICATIONS 2,397 CITATIONS
SEE PROFILE
SEE PROFILE

Some of the authors of this publication are also working on these related projects:

A global meta-analysis on the relationship between the size of outplanted seedlings and field survival View project

Forest Nursery Pathogens View project

All content following this page was uploaded by R. Kasten Dumroese on 27 February 2015.

The user has requested enhancement of the downloaded file.

 Seed Germination and
Sowing Options 9
Tara Luna, Kim M. Wilkinson, and R. Kasten Dumroese

Seeds of many native species are challenging to germinate. One important thing a
grower can do is to learn as much as possible about the life history, ecology, and habi-
tat of the species he or she wishes to grow to understand the processes seeds from each
target species go through in nature. Any observations will be valuable when trying
to germinate and grow species that have little or no published information available.
How seeds are handled, treated, and sown can affect the genetic diversity and the qual-
ity of the crop produced. Growers need to balance the desire for uniform crops and
schedules with the need to protect the diverse characteristics within species. In this
chapter, we discuss seed characteristics, treatments to improve or stimulate germina-
tion, and different types of sowing options for seeds.

Facing Page: Hand-sowing. Photo by Tara Luna.

163
Seed Characteristics Tropical species inhabiting areas with a strong wet-dry
seasonal cycle, arid or semiarid climates, or at high eleva-
As discussed in Chapter 8, Collecting, Processing, and
tions subjected to cold temperatures often have dormant
Storing Seeds (and shown in figure 8.8), tropical seeds can be
seeds. The degree of dormancy in these species can vary
divided into four categories related to their longevity and abil-
among and within seed lots, between seed crop years,
ity to be stored (Hong and Ellis 2002, Kettle and others 2011).
and individuals. Examples of species with dormant seeds
Viviparous: seeds that germinate before they are dis- include acacias such as Hawai‘i’s native Acacia koa, and
persed from the mother plant. The most common examples pines including the Caribbean’s Pinus caribaea.
are some species of mangroves such as Avicennia and
Dormancy may be caused by factors outside (external)
Rhizophora and some tropical legumes.
or inside (internal) to the seeds. Some species have a com-
Recalcitrant: seeds that germinate soon after matura- bination of external and internal dormancy, a condition
tion and dispersal from the mother plant, and cannot be known as double dormancy. Knowing the type of seed dor-
dried without losing viability. Most species from the wet, mancy is essential for successful propagation.
humid tropics have recalcitrant seeds because conditions
in these environments are consistently favorable for germi- External Seed Dormancy
nation and seedling establishment. Examples of common
species with recalcitrant seeds include cacao, mango, lon- External seed dormancy may be physical, physical-
gon, and jackfruit. physiological, chemical, or mechanical (Baskin and Baskin
1998, 2004). Seeds that have hard, thick seedcoats that physi-
Intermediate: seeds that can germinate immediately cally prevent water or oxygen movement into seeds have
but may also survive partial drying without losing viability. physical dormancy. Physical dormancy is the most common
For example, papaya seeds that have been dried to 10-per- seed dormancy type seen in the tropics. Species with external
cent moisture content have been stored successfully under dormancy include many of the legumes (Fabaceae), mallows
conditions of 50-percent relative humidity for 6 years (Malvaceae), and other tropical species that are adapted
without affecting viability (Vozzo 2002). Species that have to fire, or inhabit arid to semiarid island habitats or areas
shown intermediate storage behavior include neem, cinna- with pronounced wet-dry seasonal cycles. These seeds nor-
mon, citrus, mahogany, and coffee. mally germinate over a period of several years. Depending
Orthodox: seeds that can be dried without losing via- on species and habitat, various environmental factors cause
bility. These seeds are considered “dormant” and often these seeds to become permeable over time or during a cer-
require specific treatments to encourage germination. Dor- tain time of year. Seeds that require additional exposure to
mant seeds will not germinate immediately upon matura- particular temperatures after they become permeable have
tion and dispersal from the mother plant even when ideal physical-physiological dormancy.
environmental conditions exist. The fruits that enclose the seeds cause other forms of
Before attempting to grow a plant, it is important to external dormancy. Chemical dormancy describes fruits
know the seed germination type because that helps deter- that contain high concentrations of germination inhibitors
mine the best seed treatments and sowing options for that that prevent spontaneous germination of seeds. Mechanical
seed. For orthodox seeds, knowing about the species helps dormancy describes tough, woody fruit walls that restrict
you to provide the best conditions to dissipate, or “break,” seed germination and is best exemplified by the husks that
seed dormancy and achieve good rates of germination. surround coconuts (Cocos nucifera).

Dormancy in Orthodox Seeds Internal Seed Dormancy

Dormancy is an adaptation that ensures seeds will ger- Internal dormancy may be morphological, physiological,
minate only when environmental conditions are favor- or both (Baskin and Baskin 1998). Seeds with morphological
able for survival. The conditions necessary to allow seeds dormancy have an underdeveloped embryo when dispersed
to break dormancy and germinate can be highly variable from the mother plant. A period of after-ripening (usually
among species, within a species, or among seed sources warm and moist conditions) is needed for the embryo to fully
of the same species. This degree of variability is advanta- mature before the seed is capable of germination. Tropical
geous because seeds will germinate at different times over a species that exhibit morphological dormancy are found in the
period of days, weeks, months, or even years, ensuring that Annonaceae, Dilleniaceae, Magnoliaceae, and Myristicaceae
some offspring will be exposed to favorable environmental families and in many palm species. Seeds of this type may not
conditions for survival. germinate for several months to 1 year after sowing.

164 Tropical Nursery Manual

 Physiological dormancy is found in some species
in arid and semi-arid tropical environments. Seeds are
permeable to water, but certain environmental condi-
tions are necessary to modify the internal chemistry of
the seed and thus enable germination. Usually a period
of cold, moist conditions or holding seeds in dry storage
overcomes physiological dormancy.
Seeds with morphological-physiological dormancy
usually require a combination of warm and cold condi-
tions, often over an extended period of time, before they
are capable of germination.

Treatments To Overcome Figure 9.2—Seeds that are not cleaned before treatment or
Seed Dormancy and Enhance sowing can easily mold or be susceptible to serious pathogens
Germination such as damping-off disease. Photo by Thomas D. Landis.
A variety of seed treatments have been developed in sections. Nondormant seeds are planted immediately after
response to the diversity of seed types grown in nurseries. collection and cleaning. Intermediate seeds may be stored
Before treating seeds, be sure to consult available references for several weeks or months in suitable conditions, and then
to see what treatments have been used on that species; see cleaned, fully rehydrated, and sown. For dormant (ortho-
the literature cited at the end of this chapter and the Native dox) seeds, dormancy must be overcome using one or more
Plant Network (http://www.native plantnetwork.org). If no of the methods described in this chapter before seeds can be
information is available, check references for closely related rehydrated, enabling germination. It is essential to determine
species. Any personal observations made on the species in the which type(s) of dormancy the seed has so you can do what
habitat may also provide some clues on how to germinate the is needed to overcome dormancy (figure 9.1). The nursery
seeds. In general, however, the process of treating seeds fol- must determine whether seeds will need to be cleaned, scari-
lows a fairly standard progression outlined in the following fied, soaked, stimulated, stratified, and treated in other ways
before sowing on a species-by-species basis. The following
sections describe the seed treatment options available.

Cleaning
Seed cleaning helps prevent diseases in the nursery. Clean-
ing seeds of bacterial and fungal infestation is especially nec-
essary for species that easily mold (figure 9.2). Often, molding
can be related to the most common disease seen in nurseries,
damping-off. Seed cleaning is especially important in humid
climates and for species that take a long time to germinate.
Often, without cleaning, seeds can be lost to pathogens before
they are planted in the nursery.
One of the best cleaning methods is to simply soak seeds
in a stream of running water for 24 to 48 hours. The run-
ning water flushes bacterial and fungal spores from the seeds
(James and Genz 1981). This treatment can also be used to
satisfy the soaking requirement described in the next section.
Seeds can also be cleaned with several chemicals, some
of which also act to stimulate germination. Bleach (5.25
percent sodium hypochlorite) is the most common chem-
ical used. Depending on the species, bleach cleaning solu-
Figure 9.1—Key to dormancy types. Knowing the type of seed tions range between one part bleach in eight parts water to
dormancy is essential to successful seed propagation. Illustration
by Jim Marin. two parts bleach in three parts water. With most species,

Seed Germination and Sowing Options 165

treatment duration is 10 minutes or less. Species with very
thin seed coats should not be cleaned with bleach. Hydro-
gen peroxide can be an effective cleanser and can some-
times enhance germination (Narimanov 2000). The usual
treatment is one part peroxide in three parts water. Tropical
species that benefit from hydrogen peroxide rinses include
Albizia species and camphor tree seeds (Vozzo 2003).

Scarification
Seeds with external dormancy require scarification.
Scarification is any method of disrupting an impermeable
seed coat so that water and oxygen can enter the seeds.
In nature, hard seed coats are cracked or softened by fire,
Figure 9.4—Hobby-size rock tumblers can be used to scarify seeds
extreme temperatures, digestive acids in the stomachs of and avoid seed destruction that can occur with sulfuric acid or heat
animals, or by the abrasion of blowing sand. After the seed scarification. Photo by Tara Luna.
coat has been disrupted, oxygen and water pass into the
seeds and germination can proceed. species is helpful. Trying several methods and recording
Seeds can be scarified many ways. How well the method the results will help determine the best method for that spe-
works depends on the species and the thickness of the seed cies and seed source.
coats. Whichever method is chosen, it is very important not
to damage the endosperm, cotyledons, or embryo during Mechanical Scarification
the treatment. Taking time to learn seed anatomy of the Mechanical scarification includes filing or nicking seeds
by hand and is most often used on large-seeded species such
as Acacia, Cassia, and Sesbania (figure 9.3). Be sure to scar-
A ify on the side of the seed opposite the embryo. It is often
done one seed at a time with a nail clipper. This method is
time consuming and requires precision to adequately scarify
the seed coat without damaging the internal portions of the
seed. Sandpaper can be used on smaller seeded species such
as sedges; placing seeds into a shallow wooden box and then
rubbing them under a block of wood covered in sandpaper is
the simplest technique. Often, however, the degree of scarifi-
cation achieved with sandpaper can be variable.
Hobby-size rock tumblers can be used to process large
batches of seed more quickly than manual mechanical
scarification (figure 9.4). Dry tumbling involves placing
seeds, a coarse carborundum grit (sold by rock tumbler
B dealers), and pea gravel in the tumbler and tumbling for
several hours or several days. Wet tumbling includes the
addition of water to the grit and pea gravel. A benefit of wet
tumbling is that seeds are soaked in well-aerated water and
chemical inhibitors may be leached from the seed.

Heat Scarification
Many species, especially those from fire-adapted eco-
Figure 9.3—Mechanical scarification works well with large, systems, respond to germination cues from heat. Using
easy-to-handle seeds. Great care is needed, however, so that the either wet or dry heat to scarify the seeds can simulate this
embryo or cotyledons are not damaged. Koa seed hand scarifica- response. Using wet heat is an effective method for many
tion (A), scarified lotus seeds (B). Photo A by Craig R. Elevitch, small-seeded species because it provides a rapid, uniform
and photo B by Cardno JFNew Native Plant Nursery. treatment that can be assessed within a few hours. Wet-

166 Tropical Nursery Manual

 duration is known, the entire seedlot can be treated. Sulfuric
acid is very dangerous to handle and requires special equip-
ment, personal protective gear, and proper disposal after use.
It should never be poured down sink drains. If acid is being
diluted in water, the acid must be added to the water, never
add water to acid—when water is added to acid, heat will
be released, risking an explosion and other dangers. Some
species have thick seed coats but can easily be damaged by
sulfuric acid. Instead, citric acid or sodium or calcium hypo-
chlorite baths with longer treatment durations may be used.
The safe use of sulfuric acid requires the following proce-
dures:
Figure 9.5—Seeds that have been scarified by hot water are • Treat seeds that are dry and at room temperature.
visibly larger than untreated seeds because the seed coat has been
breached and seeds can then absorb water increasing their size. • Require workers to wear safety equipment, includ-
Photo by Greg Morgenson. ing face shield, goggles, thick rubber gloves, and full
protective clothing.
heat treatments are effective for many tropical species • If diluting, add acid to water, never water to acid.
including Acacia, Cassia, Senna, Sesbania, and Tama-
• Immerse seeds in an acid-resistant container, such as
rindus (Vozzo 2002). Because the thickness of the seed
a glass, for the duration required.
coat may vary among sources, it is wise to dissect a few
seeds and examine the thickness of their seed coats to • Stir seeds carefully in the acid bath; a glass rod
help determine treatment duration. Seeds are added to works well.
boiling water for 5 to 10 seconds and then immediately • Immerse the container with seeds and acid in an
transferred to a vat of cold water so that they cool quickly ice bath to keep temperatures at a safe level for the
to prevent embryo damage. Seeds imbibe the cool water embryos (this temperature depends on the species;
for 1 day and are then ready for sowing or for stratifica- many do not need this step).
tion (figure 9.5). Some species cannot tolerate excessively • Remove seeds from the acid by slowly pouring the
high temperatures, so you may want to heat the water to seed-acid solution into a larger volume of cool water,
only 158 °F (70 °C) and monitor your results. ideally one in which new, fresh water is continually
Dry heat is most commonly used on fire-adapted spe- being added.
cies. Seeds are placed in an oven at temperatures ranging • Stir seeds during water rinsing to ensure all surfaces
from 175 to 250 °F (80 to 120 °C) from a few minutes to are thoroughly rinsed clean.
1 hour, depending on the species. The seed coat cracks
open in response to the heat. To avoid damaging seeds, this
treatment needs to be monitored closely.

Chemical Scarification
Sulfuric acid is most commonly used on species with very
thick seedcoats and with stony endocarps that surround the
embryo (figure 9.6). It has been used on some species of Aca-
cia, Albizia, Cassia, Leucaena, Parkinsonia, and Terminalia
(Vozzo 2002). Treatment length varies with the species and
often among seed sources, and it must be carefully monitored
because seeds can be destroyed if the treatment is too long.
A simple way to monitor the process is by removing seeds at
regular intervals and cutting them with a sharp knife. When
the seeds are still firm but can be cut fairly easily, the treat-
ment is probably sufficient. Another way is to run a pilot test
on a subsample of seeds. Again, remove some seeds periodi- Figure 9.6—Seeds that have been treated with sulfuric acid. Photo
by Nancy Shaw.
cally and evaluate how well they germinate. After the best

Seed Germination and Sowing Options 167

 species having underdeveloped embryos. In some cases,
it has been used to substitute for a warm, moist treat-
ment and to hasten embryo after-ripening. Sandalwood
is a species that has been successfully germinated using
gibberellic acid. Gibberellic acid can be purchased from
horticultural suppliers. Preferred concentrations vary,
but most nurseries use 500 to 1,000 parts per million
(ppm). High concentrations can cause seeds to germi-
nate, but the resulting seedlings may be of poor quality.
Therefore, it is best to experiment with low concentra-
tions first. The following are some guidelines for treating
seeds with gibberellic acid:
• Gibberellic acid takes a long time to dissolve. It may
need constant stirring or you may want to prepare it
Figure 9.7—Many species of dormant seeds benefit from 1 to sev- the day before use.
eral days of water soaking before sowing to fully imbibe seeds and
remove any chemical inhibitors within the seeds or on the seed coats. • Store unused solution away from direct sunlight.
Photo by Brian F. Daley.
• Cut unbleached coffee filters into squares and fold
them diagonally.
Soaking
• Place gibberellic acid solution evenly into an ice cube
After cleaning and scarification, seeds must have expo- tray.
sure to water and oxygen before germination can occur. The
standard procedure is to soak seeds in water for 1 to several • Place each folded coffee filter containing the seeds
days until they are fully hydrated (figure 9.7). Hydration into the wells of the tray so that it wicks up the solu-
can be checked by taking a sample, allowing it to dry until tion.
the seed coat is still wet but dull, not glossy, and weighing • After 24 hours, remove and either sow directly or
it. When the weight no longer increases substantially with place seeds into fresh coffee filters moistened with
additional soaking time, the seeds have absorbed sufficient distilled water for stratification.
water. Scarified seeds will be more obvious; the seeds will
enlarge drastically during the soak. Seeds that only had phys- Ethylene
ical dormancy can be immediately planted. As mentioned This gas occurs naturally in plants and is known to stimu-
previously, running water rinses are effective seed cleaning late the germination of some species. Ethylene gas is released
treatments that reduce the need for fungicides in nurseries from ethephon, a commercially available product. Ethephon,
(Dumroese and others 1990). Running water soaks also help used either alone or in combination with gibberellic acid, has
to remove any chemical inhibitors present on or within the enhanced the germination in doum palm (Mousa and others
seeds. An aquarium pump can be used to agitate the seeds to 1998) and may be used for other species inhabiting arid to
improve the cleaning effect and keep the water well aerated. semi-arid tropical and saline environments. It may inhibit
If seeds are not soaked with running water, change the water germination in other species, so consult the literature and
often (at least a couple of times each day). experiment before using operationally.

Germination Stimulators Smoke Treatments

Several chemicals are known to increase seed germina- Smoke stimulates germination in many fire-adapted
tion. These chemicals are usually applied after seeds are species; for example, species from the California chaparral,
fully hydrated. In general, only seeds with internal dormancy longleaf pine communities in Florida, or species from fire-
receive this treatment. Germination stimulators include gib- dependent ecosystems in Australia, South Africa, parts of
berellic acid, ethylene, smoke, and potassium hydroxide. South America, and the Mediterranean. Smoke especially
stimulates seeds that have thin, permeable seed coats that
Gibberellic Acid allow entry of smoke into the seeds (Keeley and Fothering-
Gibberellic acid is the most important plant hormone for ham 1998). Seeds can be treated with smoke fumigation, a
the regulation of internal seed dormancy and is often used method in which smoke is piped into a specially constructed
on seeds with complex internal dormancy and with those smoke tent containing seeds sown in trays (figure 9.8A),

168 Tropical Nursery Manual

 was very important in some species; a 3-minute difference
A in exposure resulted in seed mortality. Some fire species
did not germinate under heat or smoke treatments alone.
With some species, seed burial for 1 year or stratification
was required in addition to smoke exposure. All these
factors can have an effect on germination and should be
considered when determining whether to use smoke treat-
ments. Success with this novel treatment will require trials,
so good record keeping is critical.

Potassium Hydroxide Rinses

Potassium hydroxide has been used to stimulate ger-
mination in several native plant species. Optimum con-
B centration varies from 5.3 to 7.6 Molar for 1 to 10 minutes
depending on the species; longer soaks at higher concentra-
tions were found to be detrimental (Gao and others 1998).

Other Stimulants
Potassium nitrate, thiourea, and kinetin have been used
to stimulate germination in seeds, although the use of these
compounds with tropical native plants is lacking. You may
choose to experiment with these compounds on a limited
basis with difficult-to-germinate seeds.

Temperature and Moisture Treatments

Many seeds with internal dormancy require a moist
period at certain temperatures similar to what occurs in
the natural habitat before they germinate and grow. His-
torically, stratification was a temperate zone practice of
alternating layers of moist soil and seeds in barrels and
allowing these “strata” to be exposed to winter tempera-
tures. Nowadays, stratification is often used more generi-
Figure 9.8—Smoke treatments have been used to overcome seed cally to describe the combined use of moisture and any
dormancy and enhance germination rates for many native species temperature to overcome seed dormancy. We use the term
inhabiting fire-dependent ecosystems. A smoke tent for treating
seeds (A). Smoke water-treated seeds of angelica (B). Photo A by “stratification” to refer to only cold, moist treatments (rare
Kingsley Dixon, and photo B by Tara Luna. to use in the tropics, but we cover it anyway for nurseries
growing highland species). We use the term “warm, moist
or with smoke water. Smoke water is an aqueous solution treatment” instead of “warm, moist stratification.”
of smoke extract made by burning vegetation and piping Some native species with double internal seed dormancy
the smoke through distilled water or allowing the smoke require a combination of a warm, moist treatment for a
to infuse into a container of water. Seeds are then soaked period of time followed by stratification. Some species or
in the treated water (figure 9.8B). Conversely, growers can seedlots may require only a few days or weeks of stratifica-
experiment with commercially available smoke products tion, while others may require several months. As a general
such as liquid smoke or smoke-infused paper discs, or by rule, it is best to use the maximum recommended treatment.
adding ash to growing media. Also keep in mind that what works well at one nursery may
Many variables, such as the material used for combus- not necessarily work well at another nursery because of dif-
tion, the combustion temperature, and the duration of ferences in seed source, handling, processing, cleaning, and
exposure, will need to be determined on a species-by-spe- storage. An advantage to stratifying seeds of some species is
cies basis. Experiments performed by Keeley and Fother- that it can speed up germination and make it more uniform,
ingham (1998) found that the length of exposure to smoke which is desirable in a container nursery.

Seed Germination and Sowing Options 169

Warm, Moist Treatments stratification is preferred over natural stratification unless the
Warm, moist treatment enhances after-ripening of natural treatment provides higher rates of germination.
seeds with underdeveloped embryos. Warm, moist treated For artificial stratification of small seedlots and small
seeds are kept at temperatures of 72 to 86 °F (22 to 30 °C) seeds, seeds can be placed between sheets of moistened
for a period of time, usually in moist peat moss, sawdust, or paper towels and inserted in an opened plastic bag or sown
other substrate. Although warm, moist treatments are not on a medium in flats with drainage holes. Paper towels
commonly used on tropical species, it can be considered for need to be moist but not waterlogged, and seeds need to be
seeds with morphological or physiological seed dormancy. evenly spread across the moist paper towel to help prevent
molding (figure 9.9).
Stratification Another technique is “naked” stratification. Most conifer
Stratification (cold, moist treatment) is used on seeds with seeds, for example, are stratified in this manner (figure 9.10).
internal dormancy from temperate areas, or high-elevation Seeds are placed in mesh bags and then soaked in running
habitats in tropical regions. Some subtropical species may also water as described previously. After the seeds are hydrated,
benefit from a period of cool, moist stratification. In climates the bag is pulled from the soak, allowed to drip dry for 30 to
with four seasons, seeds sown in flats or containers in late 90 seconds, and then suspended in a plastic bag. Make sure
summer or autumn and left outdoors during winter undergo the seeds are not in contact with standing water in the bag
“natural” stratification. This technique may be preferred if the and hang the bags in the refrigerator. If naked seeds need a
species has double dormancy (requires both a warm, moist warm, moist treatment before stratification, it is easiest to
treatment or stratification), requires a very long stratification first spread the seeds onto moistened paper towels enclosed
or requires low temperatures or fluctuating temperatures for in large plastic bags. After the warm treatment, the seeds can
a long period of time. Conversely, “artificial” stratification be returned to the mesh bags for stratification. One other
involves placing seeds under refrigeration at 34 to 38 °F (1 to hint: if a particular species or seedlot has a tendency to begin
3 °C) for a period of time. Artificial stratification has several germinating during stratification, surface-dry the seed coats
advantages: (1) it allows for a routine check of seeds to ensure (seeds should be moist and dull, not shiny), and then put the
they are moist and not moldy, (2) a large number of seeds can seeds into the bag for refrigeration. The seeds need to still
be stratified in a small space, and (3) seeds or seedlots that have enough moisture for chemical processes that dissi-
begin to germinate can be removed from the treatment and pate dormancy to occur but not enough moisture to allow
planted in the nursery as they become available. Artificial for germination.

A B

Figure 9.9—Small seeds requiring only a few weeks of stratification can be stratified by moistening paper towels and holding by corner to
let excess water drain away (A) or placing seeds onto moistened towels inserted into an unopened plastic zippered bag (B). Illustrations from
Dumroese and others (1998).

170 Tropical Nursery Manual

 A B

Figure 9.10—Naked stratification: After soaking seeds in a mesh bag, allow the bag to drip dry for 1 minute and then suspend the mesh
bag in a plastic bag (A). Hang the bag in the refrigerator. Make sure the seeds are not in constant contact with standing water in the plastic
bag (B). Illustrations from Dumroese and others (1998).

Many wetland and aquatic species can be treated with need to be sown on the surface of the medium so they are
naked stratification in water. In general, these species can exposed to light during germination. Other species are
be easily stratified in Ziploc®-type bags filled with water. conditioned to germinate only if they are buried in the
Insert a soda straw into the bag, ensuring that the end is soil. Species requiring darkness to germinate are those
sticking out of the bag, to allow some oxygen to reach the that germinate readily under the deep shade of a closed
seeds. Then, seal the rest of the bag securely. Place under forest canopy. Tropical trees and shrubs with medium to
refrigeration if in need of a cold, moist stratification period. larger sized seeds often require darkness for maximum
germination, but shade tolerant vines and herbaceous
Environmental Factors plants may have smaller seeds. Other species requiring
Influencing Germination darkness to germinate include some of the species that
colonize sand dunes along coastlines.
Four environmental factors affect germination: light,
water, oxygen, and temperature. All plants have specific
germination requirements based on ecological adaptations
and the environmental cues that trigger germination for
that species.

Light
Light quality and duration can influence germination.
In nature, seeds of tropical pioneer species require high
light levels associated with a canopy gap for germina-
tion and establishment, whereas shade tolerant species
generally can germinate in very poor light or deep shade.
Many small-seeded, tropical native species fall into this
category. Thus, pioneer species, such as ‘ōhi‘a in Hawai‘i,
with very small dust-like seeds (figure 9.11), require light
for germination and fail to germinate even if they are bur- Figure 9.11—In general, small-seeded pioneer species such as
‘ōhi‘a require light to germinate and must not be buried. Photo
ied only 2 mm deep (Drake 1993). Therefore, these seeds by Tara Luna.

Seed Germination and Sowing Options 171

Water and Oxygen seeds. Many tropical tree species germinate to their high-
Water is also important for germination. Overwater- est percentages at alternating temperatures of 86/68 °F
ing seeds during germination results in reduced levels of (30/20 °C) or are provided with at least 10 °F (5.6 °C)
oxygen in the medium and promotes tissue breakdown difference between the day and night time temperature.
and disease whereas underwatering delays or prevents Some difficult-to-germinate tropical species may require
germination. Therefore, seeds need to be kept evenly even greater temperature fluctuations.
moist during germination. Although oxygen is needed
for respiratory processes in germinating seeds, some Seed Sowing Methods
aquatic species may require low oxygen levels for germi- Several sowing techniques have been used for native
nation. For example, tropical floodplain forest or wetland plants (table 9.1) and are described in the following sec-
species naturally germinate during periods of high water tions. The process of sowing seeds for nursery production
inundation and respond positively to low water oxygen will vary with the species, type of seed, seed quality, and
levels (Kurbitzky and Ziburski 1994, Vozzo 2002). nursery environment.

Temperature Direct Sowing

Temperature influences seed germination rate and Direct sowing is fast, easy, and economical because
percentage. Some germination patterns in response to it minimizes seed handling and labor. It can be mecha-
temperature include seeds that require cool tempera- nized when done on a large scale. For direct sowing to be
tures, tolerate cool temperatures, require warm tempera- efficient, the seeds need to be easy to handle, abundant
tures, and (or) require alternating temperatures (Hart- in supply, have simple dormancy treatments, and have a
man and others 1997, Vozzo 2002). Species requiring known high-germination rate (figure 9.12). If the direct
cool temperatures generally germinate below 77 °F (25 sowing will be mechanized, seeds must also be uniform in
°C), which coincides with high elevations in tropical or size and shape.
subtropical regions. Species that tolerate cool tempera-
The success of direct seeding depends on the accuracy
tures will germinate over a wide range of temperatures
of seed germination information. Growers must realize
from 41 to 86 °F (5 to 30 °C). Many species will not
that actual seedling emergence may be different from
germinate under excessively high temperatures. Most
the results of laboratory germination tests that are con-
tropical species require warm temperatures and will only
ducted under ideal environmental conditions. Nursery
germinate if temperatures are above 70 °F (21 °C). In
managers must adjust for this discrepancy based on their
addition, some species germinate better when exposed to
own operational experience. Growers should conduct a
alternating temperatures. Alternating temperatures are
small germination test of each seedlot to determine the
particularly important with dormant, freshly harvested

A C

Figure 9.12—Direct sowing works well

for seeds that have little or no dormancy
(or have been treated to overcome dor-
mancy), are easy to handle and are in
abundant supply (A). Simple tools like a
film canister (B) or a folded envelope (C)
can be used to accurately sow small seeds.
B
Photo A by Douglass F. Jacobs, and pho-
tos B and C by Dawn Thomas.

172 Tropical Nursery Manual

Table 9.1—Methods for sowing seeds. Adapted from Landis and others (1999).

Good method for

Propagation method seeds with the following Advantages Disadvantages
characteristics

Direct Sowing: • Have a known high- • Fast and easy • Less efficient use of space, seeds,
Seeds are sown into percentage germination • Economical and/or growing medium
containers • Are inexpensive • Minimizes seed handling • Causes of poor germination are
• Are in abundant supply • Seeds are all sown at once difficult to track
• Have uniform, smooth shapes • May require thinning
and/or consolidation and asso-
ciated labor costs
• Not good for large or
irregularly shaped seeds

Planting Germinants: • Are of unknown viability • Efficient use of seeds • Labor intensive
Seeds sprouting or • Are valuable or rare • Efficient use of nursery space • May result in nonuniform crop
germinating in trays or • Have unknown germination • Can adjust for unknown development
bags are sown into containers requirements seed quality or performance • Root deformation possible
while roots are just beginning
to emerge • Germinate during an extended • Requires frequent, skilled
period of time or during strati- monitoring
fication

Transplanting emergents: • Are being tested but will not be • Useful with fibrous • Not recommended for
Seeds are sown into flats or transplanted to produce a crop rooted species woody and/or taprooted
seedbeds for germination; once • Do not respond well to other • Efficient use of seeds species because of problems
germinated and leaves appear, sowing methods • Efficient use of nursery space with transplant shock and
seedlings are transplanted to • Have long or unknown or root deformation
containers • Can adjust for unknown seed
dormancy quality or performance • Requires skilled labor
• Good for trials to observe seed
performance

Miniplug transplants: • Are of unknown quality • Efficient use of space • Requires two sets of
Seeds are sown directly • Are valuable or rare • Uniform crop development containers
into small containers. • Have unknown germination • Low risk of transplant injury • Timing is critical
After germination, they are requirements • Transplanting by hand
transplanted into larger is labor intensive
containers • Have very tiny seeds
• Will be transplanted into
large containers

percentage of germination for each seedlot. Those per- optimal growth and development; if so, make sure
centages can then be used to determine the number of seeds are sown in the correct orientation.
seeds to direct sow (see table 9.2). Follow these steps for • Depending on the light requirements of the species,
successful direct sowing: cover seeds with the correct amount of mulch.
• Determine how many seeds must germinate to obtain • Gently water the seeds with a fine watering head to
the production target. press them into the growing media.
• Determine if seeds can be single-sown or will require
multiple seeds to reach the production target (see Multiple-Seed Sowing and Thinning
following sections). Sowing more than one seed into each container
• Cleanse and treat seeds as necessary to break dormancy. with the expectation that at least one will germinate is
• Sow seeds, ideally centering the seeds in each con- the most common direct-sowing practice. The number
tainer. Some seeds require a specific orientation for of seeds to sow can be calculated based on the seeds’

Seed Germination and Sowing Options 173

 expected germination percentage. Two to five seeds are
typically sown per container. As a general rule, seeds
with less than a 50-percent germination are not recom-
mended for direct sowing because the high density of
nonviable seeds in the container may cause disease prob-
lems, more containers will need to be thinned, and many
plants will be wasted (figure 9.13). Table 9.2 provides
general recommendations for the number of seeds to
sow per container based on the germination percentage.
At some point, adding more seeds per container does not
really increase the number of containers with plants (table
9.3) but does drastically increase the number of contain-
ers with too many plants and the amount of seed wasted.
Sometimes it may be better to single-sow a few containers
than thin extra seedlings from many containers. For exam-
ple, sowing a single seed per container of a seedlot with
Figure 9.13—Calculating and testing germination rates help
85-percent germination yields 15 percent empty containers
reduce costs and problems associated with thinning. Photo by whereas sowing two seeds per container yields only 2 per-
Thomas D. Landis. cent empty containers, but sowing the extra seed requires

Table 9.2—For a given seed germination, increasing the number of seeds sown per container increases the number of filled
containers. In general, a target of 90- to 95-percent filled containers is reasonable. Adapted from Dumroese and others (1998).

Percentage of containers with

Seed germination percentage Seeds to sow per container
at least one seedling

90 + 1 to 2 90 to 100

80 to 89 2 96 to 99

70 to 79 2 91 to 96

60 to 69 3 94 to 97

50 to 59 4 94 to 97

40 to 49 5 92 to 97

Table 9.3—A sowing example for a seedlot of Acacia koa having a 65-percent germination rate. Assuming 1,000 seedlings are
desired, notice that adding more than three seeds per container really does not improve the number of containers with seedlings and
wastes many seeds. Adapted from Dumroese and others (1998).

Seeds sown Containers with at Seed Seedlings

Empty containers (%)
per container least one seedling (%) sown produced

1 35 65 1,000 650

2 12 88 2,000 880

3 4 96 3,000 960

4 1 99 4,000 990

5 0 100 5,000 1,000

174 Tropical Nursery Manual

thinning 72 percent of the containers. The nursery manager Oversowing works best if the nursery has extra space and
may have been better off, in terms of seed use efficiency and is using containers with individual, exchangeable cells because
labor, to have simply oversown 10 percent more containers containers with live plants can be consolidated and the extra
rather than pay for the labor to thin. Therefore, the amount containers can be removed (see Chapter 7, Containers). Single
of seeds to sow per container is a function of germination, sowing is efficient because no seeds are wasted and plants that
seed availability, nursery space, thinning costs, and so on. do emerge are not subjected to competition or the stresses of
When more than one seedling germinates in the same thinning as they are with the multiple-sowing technique. Over-
container, seedlings compete for light, water, and nutri- sowing wastes potting materials and bench space, however,
ents. This competition results in lower initial growth rates and consolidating the empty containers is labor-intensive.
and requires that seedlings be thinned (clipped, culled, or
removed from the container). For this reason, thinning Sowing Germinants
should be done as soon as possible after seedlings emerge. Germinant sowing (“sowing sprouts”) is the practice
Thinning is a labor-intensive practice and it can damage of sowing seeds that are germinating (or sprouting) into
remaining seedlings if done improperly. Train workers to the container when their young root emerges (figure
thin plants carefully and to follow these guidelines: 9.14). When done properly, germinant sowing ensures
• Thin germinants as soon as possible; the more developed that one viable seed is placed in each container, thereby
the root system becomes, the more difficult it is to thin. making efficient use of space and seeds. The resulting
seedlings are often larger because they can begin to grow
• Retain the strongest seedling closest to the center of
immediately without competition. This technique can be
the container. Thinning is an opportunity for select-
labor-intensive but results in minimal waste of materials
ing the healthiest seedling while removing inferior
and space. Sowing germinants works best for seeds that:
plants.
• Are from a rare or valuable seedlot.
• Pull or cut extra plants. For species with a long, slen-
der taproot at germination (such as pine seedlings), • Have a low or unknown germination percentage.
extra seedlings can be easily pulled before they develop • Are large or irregularly shaped.
secondary roots. For species with vigorous, fibrous root • Germinate in stratification.
systems, cutting extra plants at the stem with sharp
scissors or nipping them with fingernails may be better. • Have deep dormancy and germinate over a long
period of time.
• Discard culled plants into compost or waste.
• Rapidly produce a long root after germination (such
• Check the remaining seedling and correct any disrup- as many desert and semidesert species).
tions caused by the thinning process. (For example, if
thinning disrupted the mulch, adjust it so the seedling Germinant sowing is a relatively simple process. Seeds
has the best environment possible.) are treated as necessary, then germinated in trays or bags.

Single-Seed Sowing
Sometimes, particularly when seeds are scarce or costly or
are expected to have close to 100-percent germination, sin-
gle seeds can be directly sown into containers. This practice
ensures that every seed has the potential to become a plant
and no thinning will be necessary. If a particular number of
plants are required, then extra containers are planted, often
referred to as “oversowing,” to make up for any empty cells.
The number of extra containers to sow can be calculated
based on the percentage of germination. If a seedlot has only a
78-percent germination, for 100 plants, you must sow at least
28 extra containers (100 desired seedlings/0.78 success rate =
128 containers required). The number of oversown contain-
ers may need to be increased to account for seedling losses
during the growing cycle.
Figure 9.14—Germinants must be sown as soon as the radicle
emerges from the seed coat. Photo by Tara Luna.

Seed Germination and Sowing Options 175

 A

Figure 9.16—For transplanting emergents, seeds are hand-

B sown in trays that are usually filled with about 2 in (5 cm) of
peat moss-vermiculite growing medium. To prevent root defor-
mities, it is important the trays be deep enough that the roots
will not touch the bottom before the emergents can be trans-
planted. Photo by Thomas D. Landis.

clean scissors. The germinating seed is carefully placed in

the container with the radicle extending downward. After
the seeds are properly planted, the medium needs to be
firmed around the root and the seed covered with mulch.
An advantage of planting germinants is that the germi-
nation process is more visible to the growers than when
Figure 9.15—When planting germinants, seeds must be sown seeds are direct sown. Germination timing can be better
as soon as the radicle is visible and must be oriented correctly monitored and the causes of germination problems are eas-
when planted. Incorrect orientation leads to severe root deforma- ier to track, but because seeds in trays or bags are very close
tion in woody species (A, B). Photo A by Thomas D. Landis, and together, a mold or pathogen can contaminate all the seeds
photo B by R. Kasten Dumroese. if not properly monitored. Labor is required to routinely
check for germination, skill is required to achieve proper
Seeds may be spread out between layers of moist paper tow- planting orientation of the seeds, and planting must be
els or moist cardboard. Larger seeds are sometimes placed done in a timely fashion. Because germinants may emerge
in plastic bags filled with a moist medium such as Sphag- over several weeks or longer for some species, crop devel-
num peat moss. Seeds are closely spaced, but far enough opment will be more variable and require special cultural
apart so that mold does not spread if it forms. Seeds are treatments.
checked every few days. After seeds begin to germinate,
they must be checked daily. Germinated seeds are removed Transplanting Emergents
daily and planted directly into growing medium in their
containers. Larger seeds can be planted by hand; smaller Transplanting emergents (“pricking out”) is a practice
seeds are often sown using tweezers. for germinating seeds in a small area. Seeds are hand-sown
in shallow trays that are usually filled with about 2 in (5 cm)
Timing and root orientation are critical when sowing of peat moss-vermiculite (or similar) growing medium (fig-
germinants. Seeds need to be sown into containers as soon ure 9.16). Soon after the seeds germinate, they are “pricked
as the root emerges. The embryonic root, often called a out” of the tray and transplanted into a container. This
“radicle,” needs to be short, ideally no longer than 0.4 in (1 technique is not recommended for woody plants and other
cm). If the radicle becomes too long, it may be difficult to taprooted species, because root problems often result.
plant without causing root deformation (figure 9.15). Some
growers like to prune the radicle of taprooted species before Transplanting emergents works best when—
planting to ensure a more fibrous root system. No more • Species have a fibrous root system that recovers well
than the very tip (up to 0.1 in [3 mm]) is trimmed with from transplanting (herbaceous forbs without a tap-
root, grasses, sedges, and rushes).

176 Tropical Nursery Manual

 • Tests or trials are being used to observe seed treat-
ments, germination timing or percentage, early
growth rate or other early developmental issues.
• Seeds are too small or fragile to be sown by any other
method.
• Seeds have very complex dormancy or germinate over
an extended period of time.
• Limited nursery growing space makes direct seeding
uneconomical.
• Timing to transplant emergents is scheduled
promptly.
Figure 9.17—Root and shoot development at various stages
Some key disadvantages include the following— after germination of the Caribbean coastal shrub, Chrysobala-
• Disease potential is high in densely planted trays. nus icaco. The large-seeded species is easily transplanted at
these stages. Photo by Brian F. Daley.
• Root orientation and timing is critical; root malforma-
tions and other problems can result if neglected.
favorable environment for germination. Although the exact
• Transplanting is skill and labor intensive. size or age to transplant the germinating seedlings varies
Training, care, good timing, and proper technique is by species, it is usually done at the primary leaf stage (after
required to prepare seedling trays, sow the seeds, and to cotyledons emerge) and well before root systems reach the
transplant emergents properly. As with many other nursery bottom of the seed tray (figure 9.17).
functions, some trial and error occurs in finding the medium Emergents are carefully removed from the tray, usually
mixture and tray depth that works best for each species. by gently loosening the medium around them (figure 9.18A).
Larger seeds are scattered by hand over the surface of the A small hole is made in the medium of the container and the
moistened medium, or place in indentations in the medium. germinant is carefully transplanted, ensuring proper root
Smaller seeds can be sown with a salt shaker with enlarged orientation (figure 9.18B). Some species benefit from
holes. Sown seeds are then covered with a light application root pruning before transplanting. The potting medium
of fine-textured mulch or medium, irrigated, and placed in a is then firmed around the root and stem (figure 9.18C).

A B C

Figure 9.18—Transplanting emergents works well for fibrous-rooted shrubs, forbs, and grasses. Great care must be taken to lift
the emergent from the pricking out tray without damaging the roots (A) and to carefully and properly transplant it into the new
container filled with moistened growing media (B, C). Photos by Tara Luna.

Seed Germination and Sowing Options 177

 When timed incorrectly or done improperly, especially
on taprooted woody species, transplanting emergents can
produce a “J-root” or kink in the seedling stem or root
(figure 9.19). These malformations can cause mechanical
weakness, poor growth in the nursery and later in the field,
and mortality after outplanting. Therefore, unless no other
sowing method works, transplanting emergents of woody
plants is discouraged.

Transplanting Plugs
Small-volume containers, such as miniplugs (figure
9.20) or expanded peat pellets, in which seeds are direct
sown (see Chapter 7, Containers) can be transplanted
into a larger container after the seedlings are well estab-
lished. Transplanting small plugs has a number of ben- Figure 9.20—Miniplugs are a viable option for growing seed-
efits. The small plug container preserves healthy root lings that will later be transplanted to a larger container. Mini-
form because damage to roots during transplanting is plugs work very well with species with very tiny seeds. Photo by
eliminated. Planting small plugs also makes efficient use Tara Luna.
of growing space. Large numbers of small plugs can be
started in a very small area and managed intensively dur- transplanting process, but they must not have so many
ing germination and early growth. roots that they are rootbound or the roots may become
Plants in miniplug containers must have a firm enough deformed after transplanting. If peat pellets are used, too
root plug to hold the plug together and withstand the few roots are not a problem because the entire pellet can
be transplanted. A hole large enough to accept the plug
is made in the medium of the larger container, and the
small plug-grown seedling is carefully inserted. Planters
need to ensure that the roots go straight down and are
not deformed during transplanting. The medium is gen-
tly firmed around the root system, mulch is applied, and
the plant is watered.
Transplanting small plugs is labor intensive and
requires skill. In some arid, windy areas, small plugs are
not practical because they dry out too quickly between
waterings. Before investing in small plugs on a large scale,
a small trial is advised.

Seed Coverings (Mulch)

Regardless of the seed sowing method, a seed cover
or “mulch” is recommended to create an optimal envi-
ronment for germinating seeds. The only exception is for
species that require light to germinate. Mulch is usually a
light-colored, nonorganic material spread thinly over the
seeds. Examples of mulches include granite grit (such as
poultry grit) (figures 9.21A, 9.21B), pumice, perlite (fig-
ure 9.21C), coarse sand, or vermiculite (figure 9.21D).
When properly applied, mulches—
Figure 9.19—Transplant emergents early to avoid root malfor- • Create an ideal “moist but not saturated” environ-
mation. This palm seedling was permitted to get too large in the
tray, causing the root to grow at a 90° angle. This root malforma- ment around germinating seeds by making a break
tion will cause poor growth and performance later in the plant’s in the texture of the potting medium (water will not
life. Photo by Brian F. Daley. move from the medium into the mulch).

178 Tropical Nursery Manual

 A C

Figure 9.21—Seed mulches are im-

portant to hold the seeds in place and to
moderate the surface temperature of the
medium during germination. Common
mulches include poultry grit (A, B) per-
lite (C), and vermiculite (D). Photos A
and B by Craig R. Elevitch, and photos C B D
and D by Thomas D. Landis.

• Keep seeds in place. This practice improves contact a fine-textured material such as fine-grade perlite or milled
with the medium and minimizes the number of seeds Sphagnum peat moss. Uncovered and barely covered seeds
washed out of the containers by irrigation or rainfall. must be misted frequently to prevent them from drying
• Reflect heat when mulches are light colored, so seeds out. After light-requiring and light-sensitive species have
do not get too hot on bright, sunny days. emerged and are well established, mulch can be applied to
prevent moss and liverwort growth and to help keep the
• Reduce the development of moss, algae, and liver-
medium moist.
worts (figure 9.22).
The recommended depth of the seed covering varies by
species; a general rule is to cover the seed twice as deep as
the seed is wide. If mulch is too shallow, seeds may float
away in the irrigation water. If the mulch is too deep, small
plants may not be able to emerge above it (figure 9.23).
Seeds requiring light need to be left uncovered. Very
small seeds need to be left uncovered or barely covered with

Figure 9.23—A general rule of thumb for covering seeds with

mulch is to cover the seed twice as deep as the seed is wide. Spe-
cies requiring light for germination should never be covered
Figure 9.22—Mulches help to prevent the development of with mulch, although mulch can be added after germination to
mosses and liverworts, which can compete with the seedling. reduce the growth of moss, liverworts, and weeds. Illustration
Photo by Thomas D. Landis. by Jim Marin.

Seed Germination and Sowing Options 179

 Germinating Fern Spores
Ferns (figure 9.24) and fern allies (moonworts, mosses, and horsetails) differ
from seed plants in that they produce spores instead of seeds. An understanding of
the life cycle of ferns is essential for successful fern propagation in nurseries.
Ferns have two life stages—the gametophyte and the sporophyte—the latter being
the spore-producing fern plant with which we are all familiar. The sporangia (spore-
bearing structures) are variously placed on the lower surface of the leaves and occur in
clusters known as sori. In many species, the sori are covered by specialized outgrowths
of the leaf, known as the indusium, which lifts and shrivels when spores are ripe. A
specialized layer of cells on the stalks of the spores, known as the annuli, contract and
expand and the mature spores are disseminated with a catapult-like discharge.
After the spores disseminate, they germinate upon contact with a suitably moist Figure 9.24—Native Hawaiian fern, a Sadleria
substrate. Spore germination results in the gametophyte, which begins development species. Photo by J.B. Friday.
as a small, pale-green, algae-like chain of cells known as the germ filament. Develop-
ment continues into a flat, heart-shaped structure called the prothallus. Slender holdfasts, known as rhizoids, develop on the lower surface
of the prothallus. The reproductive structures, the antheridia (male) and the archegonia (female), develop on the lower surface of the
prothallus. Antheridia usually appear before the archegonia, mostly near the rhizoids. Archegonia appear near the notch of the prothallus.
Water must be present for the sperm to swim from the antheridia to the eggs in the archegonia. After fertilization, the young sporophyte
receives its nutrients from the gametophyte via a foot-like structure. Further development is rapid and, after the sporophyte achieves a level
of photosynthesis sufficient to maintain itself, the gametophyte disintegrates. The sporophyte completes the life cycle when it grows into a
mature fern plant and produces spores.
Fern Propagation
When propagating ferns, maintaining a high level of sanitation during all phases of fern development is of the utmost importance. To
collect fern spores, find fern fronds with ripe (dark brown, black, or gold) sori. Cut a piece off the frond. Lay each frond piece in a paper enve-
lope, paper bag, or between two sheets of paper. Wait 24 to 48 hours for the spores to drop onto the paper, then collect (Lilleeng-Rosenberger
2005). Removal of nonspore material reduces the chances of contamination by fungi, algae, or bacteria during spore germination. Spores
can be sterilized by adding them to a 2- to 5-percent bleach solution with a small drop of wetting agent, which prevents the spores from
sticking together. Soak for a few seconds to 1 minute. Collect spores on filter paper and rinse with distilled water thoroughly for 2 minutes
(Hoshizaki and Moran 2001).
Germinate spores using sterilized propagation flats that are at least 2 in (5.2 cm) deep with drainage holes. You can use peat cubes, peat
pellets, or a sterilized clay brick placed within an airtight, clear plastic shoebox container. Any sterilized commercial growing mix without
added fertilizers can also be used. Moisten the medium thoroughly with distilled water before sowing.
Spores can be sown by spraying them on the medium surface with an atomizer, or by delivering small amounts of spores through a
syringe with distilled water. Another option is dip a cotton swab into an envelope filled with spores, and then apply the spores to the growing
media using the cotton swab.
After sowing, irrigate with distilled water and cover immediately with a clear plastic lid to seal in moisture and to prevent fungal con-
tamination. Place flats under 150- to 500-foot candles of light. If you are using artificial light (cool, white fluorescent), leave lights on for 8 to
24 hours per day. Optimum spore germination temperature is 68 to 86 °F (20 to 28 °C). Try to maintain a constant germination temperature
to avoid excessive condensation in the sealed flats. Distilled water needs to be applied when the medium begins to dry slightly on the surface.
Flats also need to be closely monitored for any fungal contamination. Spores germinate 10 to 20 days after sowing. If additional irrigation is
needed, water only with distilled water delivered through a sterile mist bottle or by placing flats in trays filled with distilled water.
The presence of mold may require special treatments. Stop overhead watering. Make sure that water is not dripping excessively onto
plants from condensation. Remove mold and at least 12 mm (0.5 in) of plant tissue and medium beyond the infected area. Apply a mild
fungicide labeled for ferns if infection continues.
Shortly after spore germination, the thread-like germ filament can be seen with the aid of a microscope. In general, prothalli become
visible 20 days after sowing. The prothalli continue to grow for up to 10 weeks before the reproductive structures, the antheridia and arche-
gonia, become evident on the under surface of the prothallus. These structures can be seen with a microscope when sampling a few prothalli
from a tray. After these structures appear, it is important to maintain a thin film of distilled water over the surface of the prothalli. It is very
important to keep the germination surfaces evenly moist at all times.
The clear plastic lid is removed from the container when the antheridia have withered and disappeared, usually 4 weeks after their
initial appearance. Flats are then transferred from under indoor lights to a shaded greenhouse. The young fern plants (sporophytes) with
true leaves and a developing root system will appear sometime after fertilization; from a few weeks to a few months. They can then be
transplanted into individual containers.

180 Tropical Nursery Manual

Try Different Sowing Techniques
and Keep Detailed Records
Native plant growers often work with seeds of spe-
cies that have not yet been propagated in nurseries. Little
literature or experience is available to answer questions
about seed dormancy-breaking requirements, environ-
mental requirements, germination percentages, and
other factors. Understanding the biology and ecology of
tropical plants will provide important clues on how to
overcome seed dormancy (if any) and provide the cor-
rect environmental conditions needed for germination.
It is important to develop a good recordkeeping sys-
tem to refine and improve results over time and prevent
the loss of valuable information. Keep details on the gen-
eral information of the species, seedlot, seed treatments,
and resulting germination. To improve propagation
results, use these details to develop and refine propaga- Figure 9.25—Small trials of seed germination and sowing
tion protocols, as described in Chapter 4, Crop Planning: options will help you discover the most effective approach for different
Propagation Protocols, Schedules, and Records. species. Photo by Brian F. Daley.

Trials To Develop a Successful Propagation Protocol

A for an Endangered Species
Catesbaea melanocarpa of Puerto Rico and the U.S. Virgin Islands is considered
federally endangered throughout its range. The seeds (figure 9.26A) are small and chal-
lenging to work with.
A few individuals of Catesbaea melanocarpa were discovered in a pasture after
B a wildfire burned grass and underbrush that was obscuring them. With so few plants
remaining and in danger of future fires, propagation by seed became a priority. After
months of monitoring, a dozen or so ripe fruit were collected, each containing an aver-
age of eight small, flat seeds. The seeds were too small to be sown in traditional seed
trays and monitored effectively. Due to their small size and scarcity, researchers at the
University of the Virgin Islands decided to treat the seeds similar to the way they handle
orchid seeds. Sterile dishes of sucrose rich agar (gel) were prepared and 6-8 seeds were
placed in each of 10 dishes (figure 9.26B). This intensive germination method allowed
for daily observations on every seed collected. Less than one-half of the seeds germi-
nated, and later dissection revealed the nongerminated seeds were smaller than the ger-
minated seed and did not have embryos. When the germinants produced adult leaves,
the entire dish was transplanted into plant pots with a mix of ProMix (peat, perlite, and
C vermiculite blend) and coarse sand. More than 25 percent of the plants desiccated and
died shortly after transplanting. The rest of the plants established, but grew slowly. Even
with the addition of sand, the media seemed too coarse for the tiny plants.
Later in the season, more fruit were collected. The researchers now knew that
one-half of the seeds were likely viable, but that the smaller, malformed seeds could
be discarded because they do not have embryos. Researchers added vermiculite to the
growing medium mix and sifted it through a screen. Seeds were sown directly into the
Figure 9.26—Seeds of Catesbaea melano- new, fine medium and germinated in trays in the greenhouse with 80 percent success
carpa shown here with a fruit, a fruit in cross (figure 9.26C). The second set of seedlings grew more vigorously and did not suffer the
section, and with cleansed seeds (A) were evalu- transplant shock that killed plants in the first trial.
ated in sterile dishes (B) to develop a successful The first trial led to discoveries about the seeds’ viability and identified problems
germination protocol for this endangered species with the growing medium and transplant shock. The second trial used this infor-
(C). Photos by Brian F. Daley. mation and resulted in high germination rates and healthy seedlings of a federally
endangered plant in the greenhouse.

Seed Germination and Sowing Options 181

 Because growers have a number of options for sow- In tropical ecosystems, however, recalcitrant species are as
ing seeds, it is a good idea to do small trials of several of numerous and as important as orthodox species. Because of
the methods described in this chapter (figure 9.25). See this distinction, many tropical species cannot be “banked”
Chapter 20, Discovering Ways to Improve Nursery Prac- in conventional seed banks. Instead, nurseries and seedling
tices and Plant Quality, for proper ways of conducting propagation efforts will play key roles in any efforts to con-
trials. Although several methods may “work”—that is, serve and restore tropical recalcitrant species (Kettle and
result in a viable plant produced—the question during
others 2011).
the trials should be: Which method is optimal? Trials will
help you decide how to answer this question yourself.
References
Larger Role of Nurseries in Baskin, C.C.; Baskin, J.M. 1998. Seeds: ecology, biogeography and
evolution in dormancy and germination. San Diego, CA: Aca-
Conservation of Species demic Press. 666 p.
Each seed is a link between the evolutionary processes
of the past and the potential for future adaptation (Flores Baskin, C.C.; Baskin, J.M. 2004. Determining dormancy-breaking
2002). While you clean, treat, and germinate seeds to grow and germination requirements from the fewest numbers of seeds.
your crops, be mindful how your actions affect that spe- In: Guerrant, E.O., Jr.; Havens, K.; Maunder, M., eds. Ex situ plant
conservation: supporting species survival in the wild. Washington,
cies genetic diversity and ability to adapt to the future. Do
DC: Island Press: 162–179.
your best to maintain as much genetic diversity as possible
within the species you propagate. Drake, D.R. 1993. Germination requirements of Metrosideros poly-
Nurseries are often involved in genetic selection beyond morpha, the dominant tree of Hawaiian lava flow and rainforests.
seed collection practices, intentionally or not. For example, Biotropica. 25: 461–467.
production schedules may cause growers to favor faster ger-
Dumroese, R.K.; Landis, T.D.; Wenny, D.L. 1998. Raising forest
minating over slower germinating individuals of the same
tree seedlings at home: simple methods for growing conifers of
species, although the quality of the resulting plant would
the Pacific Northwest from seeds. Contribution 860. Moscow, ID:
be similar. For some species, the earliest sprouters may be Idaho Forest, Wildlife and Range Experiment Station. 56 p.
the healthiest. But in other cases, sprouting later may be an
adaptive trait; one that could be selected out accidentally Dumroese, R.K.; Wenny, D.L.; Quick, K.E. 1990. Reducing pes-
by nursery practices. Following up with plant performance ticide use without reducing yield. Tree Planters’ Notes. 41(4):
in the field can reveal if, in fact, the slower germinating 28–32.
individuals grow well. If so, no reason exists to select these
individuals out with nursery practices, and every reason to Flores, E.M. 2002. Seed biology. In: Vozzo, J.A., ed. 2002. The
keep their traits in the gene pool. In this example, simply tropical tree seed manual. Agriculture Handbook 721. Washing-
ton, DC: U.S. Department of Agriculture, Forest Service. 13–118.
planting the slower sprouting individuals, as well as the fast
Chapter 1.
ones, could protect diversity. This example is just one that
shows how different steps in collecting, storing, germinat- Gao, Y.P.; Zheng, G.H.; Gusta, L.V. 1998. Potassium hydroxide
ing, and sowing seeds can affect subsequent plant genetics. improves seed germination and emergence in five native plant
The desire for uniform crop size and standardized schedule species. HortScience. 33: 274-276.
must be balanced with the need to protect and perpetuate
species and genetic diversity. Hartman, H.T.; Kester, D.E.; Davies, F.T.; Geneve, R.L. 1997.
Plant propagation: principles and practices. 5th ed. Upper Saddle
Nurseries may also have an increasing role in the con- River, NJ: Prentice Hall Press. 770 p.
servation of tropical species by helping protect and restore
genetic diversity of recalcitrant species (species whose Hong, T.D.; Ellis, R.H. 2002. Storage. In: Vozzo, J.A., ed. 2002.
seeds do not store well). You are probably familiar with The tropical tree seed manual. Agriculture Handbook 721.
the concept of “seed banks”—seed storage facilities used as Washington, DC: U.S. Department of Agriculture, Forest Service.
reserves to protect and restore species in case their habitats 13–118. Chapter 3.
are threatened. Seed banks are also used for some tradi-
tional food crops that have become rare with conventional Hoshizaki, B.J.; Moran, R.C. 2001. The fern growers manual.
Portland, OR: Timber Press. 604 p.
agriculture. Seed banks work well for orthodox species,
whose seeds can store for many years and remain viable.

182 Tropical Nursery Manual

 James, R.L.; Genz, D. 1981. Ponderosa pine seed treatments: Narimanov, A.A. 2000. Presowing treatment of seeds with hydro-
effects on seed germination and disease incidence. Report 81-16. gen peroxide promotes germination and development in plants.
Missoula, MT: U.S. Department of Agriculture, Forest Service, Biologia. 55: 425–428.
Northern Region, Forest Pest Management. 13 p.
Vozzo, J.A., ed. 2002. The tropical tree seed manual. U.S. Depart-
Keeley, J.E.; Fotheringham, C.J. 1998. Smoke induced seed ment of Agriculture, Forest Service. Washington, DC: U.S. Gov-
germination in California chaparral. Ecology. 79: 2320-2336. ernment Printing Office. 899 p.

Kettle, C.J.; Burslem, D.F.R.P.; Ghazoul, J. 2011. An unorthodox

approach to forest restoration. Science. 333: 35. Additional Reading
Dreesen, D. 2004. Tumbling for seed cleaning and conditioning.
Kurbitzky, K.; Ziburski, A. 1994. Seed dispersal in floodplain for-
Native Plants Journal. 5: 52–54.
ests of Amazonia. Biotropica. 26: 30–43.
Koebernik, J. 1971. Germination of palm seed. International Palm
Landis, T.D.; Tinus, R.W.; Barnett, J.P. 1999. The container tree
Society. 15: 134–137.
nursery manual: volume 6, seedling propagation. Agriculture
Handbook 674. Washington, DC: U.S. Department of Agricul-
Landis, T.D. 2000. Where’s there’s smoke…there’s germination?
ture, Forest Service. 166 p.
Native Plants Journal. 1: 25–29.
Lilleeng-Rosenberger, K. 2005. Growing Hawaii’s Native Plants.
Schmidt, L. 2007. Tropical forest seed. Berlin, Germany: Springer-
Honolulu, HI; Mutual Publishing. 420 p.
Verlag. 409 p.
Mousa, H.; Margolis, H.A.; Dubay, P.A.; Odongo, J. 1998. Factors
affecting germination of doum palm (Hyphaene thebaica Mart.)
from the semi-arid zone of Niger, West Africa. Forest Ecology
and Management 104 (1-3): 27-41.

Seed Germination and Sowing Options 183

View publication stats