germination

This article may require copy-editing for grammar, style, cohesion, tone or spelling. You can assist by editing it now.  (June 2009)

Sunflower seedlings, just three days after germination

Germination rate testing on the germination table

Germination is the process in which a seed emerges from a period of dormancy. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However, the growth of a sporeling from a spore, for example the growth of hyphae from fungal spores, is also germination. In a more general sense, germination can imply anything expanding into greater being from a small existence or germ.

Seed germination

Brassica campestris germinating seeds

A germinated seedling (Eranthis hyemalis) emerges from the ground

Germination is the growth of an embryonic plant contained within a seed; it results in the formation of the seedling. The seed of a higher plant is a small package produced in a fruit or cone after the union of male and female sex cells. Most seeds go through a period of quiescences where there is no active growth, during this time the seed can be safely transported to a new location and/or survive adverse climate conditions until it is favorable for growth. The seed contains an embryo and in most plants stored food reserves wrapped in a seed coat. Under favorable conditions, the seed begins to germinate, and the embryonic tissues resume growth, developing towards a seedling.

Requirements for seed germination

The germination of seeds is dependent on both internal and external conditions. The most important external factors include: temperature, water, oxygen and sometimes light or darkness.^[1] Different plants have seeds that require distinctive variables for successful germination. Often this depends on the individual seed variety and is closely linked to the ecological conditions of the plants' natural habitat. For some seeds, future germination responses are affected by environmental conditions during seed formation; most often these responses are types of seed dormancy.

• Water - is required for germination. Mature seeds are often extremely dry and need to take in significant amounts of water, relative to the seeds dry weight, before cellular metabolism and growth can resume. Most seeds respond best when there is enough water to moisten the seeds but not soak them. The uptake of water by seeds is called imbibition which leads to the swelling and the breaking of the seed coat. When seeds are formed, most plants store food, such as starch, proteins, or oils, to provide nourishment to the growing embryo inside the seed. When the seed imbibes water, hydrolytic enzymes are activated that break down these stored food resources in to metabolically useful chemicals, allowing the cells of the embryo to divide and grow, so the seedling can emerge from the seed.^[1] Once the seedling starts growing and the food reserves are exhausted, it requires a continuous supply of water, nutrients and light for photosynthesis, which now provides the energy needed for continued growth.

• Oxygen - is required by the germinating seed for metabolism:^[2] If a seed is in soil with too much water or is buried too deeply, it can be cut off from the necessary oxygen. Oxygen is used in aerobic respiration, the main source of the seedling's energy until it grows leaves.^[1] Some seeds have impermeable seed coats that prevent oxygen from entering the seed, causing a type of physical dormancy which is broken when the seed coat is worn away enough to allow gas exchange and water uptake from the environment.

• Temperature - affects cellular metabolic and growth rates. Different seeds germinate over a wide range of temperatures, with many preferring temperatures slightly higher than room-temperature while others germinate just above freezing and others responding to alternation in temperature between warm to cool. Some seeds germinate when the soil is cool 28-40F (-2 - 4C), while most germinate when the soil is warm 60-75F (16 - 24C) and some when the soil is even warmer 76-90F (24 - 32C). Often, seeds have a set of temperature ranges where they will germinate and will not do so above or below this range. In addition, some seeds may require exposure to cold temperature (vernalization) to break dormancy before they can germinate. As long as the seed is in its dormant state, it will not germinate even if conditions are favorable. Seeds that are dependent on temperature to end dormancy, have a type of physiological dormancy. For example, seeds requiring the cold of winter are inhibited from germinating until they experience cooler temperatures. For most seeds that require cold for germination 4C is cool enough to end dormancy, but some groups especially with in the family Ranunculaceae and others, need less than -5C. Some seeds will only germinate when temperatures reach hundreds of degrees, as during a forest fire. Without fire, they are unable to crack their seed coats; this is a type of physical dormancy.

• Light or darkness - can be a type of environmental trigger for germination in seeds and is a type of physiological dormancy. Most seeds are not affected by light or darkness, but many seeds, including species found in forest settings will not germinate until an opening in the canopy allows them to receive sufficient light for the growing seedling.^[1]

Stratification mimics natural processes that weaken the seed coat before germination. In nature, some seeds require particular conditions to germinate, such as the heat of a fire (e.g., many Australian native plants), or soaking in a body of water for a long period of time. Others have to be passed through an animal's digestive tract to weaken the seed coat and enable germination.^[1]

Malted (germinated) barley grains

Dormancy

Many live seeds are dormant, meaning they will not germinate even if the environment has sufficient water and oxygen. Dormancy is influenced by both internal and external factors that affect many different parts of the seed, from the embryo to the seed coat. Light, temperature, and pH are examples of external factors that can affect seed dormancy. Internal factors affecting seed dormancy include the presence of certain plant hormones, notably absciscic acid, which inhibits germination, and gibberelin, which ends seed dormancy. In brewing, barley seeds are treated with gibberellin to ensure uniform seed germination in the production of barley malt.^[1]

Seedling establishment

In some definitions, the appearance of the radicle marks the end of germination and the beginning of "establishment", a period that ends when the seedling has exhausted the food reserves stored in the seed. Germination and establishment as an independent organism are critical phases in the life of a plant when they are the most vulnerable to injury, disease, and water stress.^[1] The germination index can be used as an indicator of phytotoxicity in soils. The mortality between dispersal of seeds and completion of establishment can be so high that many species have adapted to produce huge numbers of seeds.

Germination rate

In agriculture and gardening, germination rate is the number of seeds of a particular plant species, variety or particular seedlot that are likely to germinate. This is usually expressed as a percentage, e.g. an 85% germination rate indicates that about 85 out of 100 seeds will probably germinate under proper conditions. Germination rate is useful in calculating seed requirements for a given area or desired number of plants.

Dicot germination

The part of the plant that emerges from the seed first is the embryonic root, termed radicle or primary root. This allows the seedling to become anchored in the ground and start absorbing water. After the root absorbs water, the embryonic shoot emerges from the seed. The shoot comprises three main parts: the cotyledons (seed leaves), the section of shoot below the cotyledons (hypocotyl), and the section of shoot above the cotyledons (epicotyl). The way the shoot emerges differs between plant groups.^[1]

Epigeous

In epigeous (or epigeal) germination, the hypocotyl elongates and forms a hook, pulling rather than pushing the cotyledons and apical meristem through the soil. Once it reaches the surface, it straightens and pulls the cotyledons and shoot tip of the growing seedlings into the air. Beans, tamarind, and papaya are examples of plant that germinate this way.^[1]

Hypogeous

Another way of germination is hypogeous (or hypogeal) where the epicotyl elongates and forms the hook. In this type of germination, the cotyledons stay underground where they eventually decompose. Peas, for example, germinate this way.^[1] Germination also starts with one tiny seedling and then begins to sprout.

Monocot germination

In monocot seeds, the embryo's radicle and cotyledon are covered by a coleorhiza and coleoptile, respectively. The coleorhiza is the first part to grow out of the seed, followed by the radicle. The coleoptile is then pushed up through the ground until it reaches the surface. There, it stops elongating and the first leaves emerge through an opening as it is.^[1]

Precocious germination

While not a class of germination, precocious germination refers to seed germination before the fruit has released to seed.^[3] The seeds of the green apple commonly germinate in this manner.^{[citation needed]}

Pollen germination

Another germination event during the life cycle of gymnosperms and flowering plants is the germination of a pollen grain after pollination. Like seeds, pollen grains are severely dehydrated before being released to facilitate their dispersal from one plant to another. They consist of a protective coat containing several cells (up to 8 in gymnosperms, 2-3 in flowering plants). One of these cells is a tube cell. Once the pollen grain lands on the stigma of a receptive flower (or a female cone in gymnosperms), it takes up water and germinates. Pollen germination is facilitated by hydration on the stigma, as well as the structure and physiology of the stigma and style.^[1] Pollen can also be induced to germinate in vitro (in a petri dish or test tube).^[4][5]

During germination, the tube cell elongates into a pollen tube. In the flower, the pollen tube then grows towards the ovule where it discharges the sperm produced in the pollen grain for fertilization. The germinated pollen grain with its two sperm cells is the mature male microgametophyte of these plants.^[1]

Self-incompatibility

Since most plants carry both male and female reproductive organs in their flowers, there is a high risk for self-pollination and thus inbreeding. Some plants use the control of pollen germination as a way to prevent this selfing. Germination and growth of the pollen tube involve molecular signaling between stigma and pollen. In self-incompatibility in plants, the stigma of certain plants can molecularly recognize pollen from the same plant and prevents it from germinating.^[6]

Spore germination

Germination can also refer to the emergence of cells from resting spores and the growth of sporeling hyphae or thalli from spores in fungi, algae, and some plants.

Conidia are the asexual reproductive spores of fungi, which germinate under specific conditions. From the germinating conidia different cells are formed. The most common one is the germ tube. The germ tube will grow and developed into the hyphae. During germination, conidial may produce conidial anastomosis tubes, those are different from conidial anastomosis tubes because they are thinner, shorter, lack branches, exhibit determinate growth, and home toward each other. Both cells have a tubular shape, but the conidial anastomosis form a bridge that allows fusion between conidia. ^[7][8]

Resting spores

In resting spores, germination involves cracking the thick cell wall of the dormant spore. For example, in zygomycetes the thick-walled zygosporangium cracks open and the zygospore inside gives rise to the emerging sporangiophore. In slime molds, germination refers to the emergence of amoeboid cells from the hardened spore. After cracking the spore coat, further development involves cell division, but not necessarily the development of a multicellular organism (for example in the free-living amoebas of slime molds).^[1]

Zoospores

In motile zoospores, germination frequently means a lack of motility and changes in cell shape, which allow the organism to become sessile.^[1]

Ferns and mosses

In plants such as bryophytes, ferns, and a few others, spores germinate into independent gametophytes. In the bryophytes (e.g. mosses and liverworts), spores germinate into protonemata, similar to fungal hyphae, from which the gametophyte grows. In ferns, the gametophytes are small, heart-shaped prothalli that can often be found underneath a spore-shedding adult plant.^[1]

See also

• Lily Seed Germination Types
• Seedling

References

External links

Wikibooks has more on the topic of Germination

• Sowing Seeds A survey of seed sowing techniques.
• Seed Germination: Theory and Practice, Norman C. Deno, 139 Lenor Dr., State College PA 16801, USA. An extensive study of the germination rates of a huge variety of seeds under different experimental conditions, including temperature variation and chemical environment.
• Starting Seeds Description of different seed germination methods.