After-ripening: period after dispersal when the seed, spores and bud cannot germinate, even under favorable conditions, and during which changes occur allowing it to germinate.
Finally, it is important to realize there is very good reason for this confusion, why seed biologists like nothing better to do than invent new terms: dormancy is an extremely complicated area of biology. No one has figured out the mechanism of embryo dormancy. Most of the literature doesn’t differentiate between dormancy imposed by the embryo and surrounding (often inhibiting) envelopes when they discuss experimental seed germination results. And then there is the problem of figuring out how these seed behave in the soil: agricultural soil seed bank dynamics.
Germinability: the capacity of an seed, bud or spore to germinate under some set of conditions.
1. A biochemical process may need to be stimulated before the germination process can begin
Dormancy: a state in which viable seeds, spores or buds fail to germinate under conditions favorable for germination and vegetative growth.
The reaction of seeds to light signals is dependent on phytochromes that consist of a group of proteins acting as sensors to changes in light conditions. Cancellation of dormancy by light is mediated by the phytochromes. All phytochromes have two mutually photoconvertible forms: Pfr (considered the active form) with maximum absorption at 730 nm and Pr with maximum absorption at 660 nm. The photoconversion of phytochrome in the red light (R)-absorbing form (Pr) to the far red light (FR)-absorbing form (Pfr), has been identified as part of the germination induction mechanism in many plant species (Gallagher and Cardina, 1998). Germination can be induced by Pfr/P as low as 10 −4 and is usually saturated by <0.03 Pfr/Pr (Benech-Arnold et al., 2000). The quality of light received by seeds may be more important than the quantity. There is evidence that Far-red light (FR, about 735 nm) can inhibit germination (Ballaré et al., 1992). Regarding the way weed emergence is influenced by light, given that FR or the ratio of FR to red light (R, about 645 nm) increases as plant canopies develop and solar elevation decreases with time after the summer period, emergence of sensitive species should be inhibited during the summer period. However, the practical significance of FR exposure for emergence in field settings is not well-known (Forcella et al., 2000).
Consequently, it is preferable to focus on depleting the seed stock in the soil through time rather than viewing weeds just as an annual threat to agricultural production (Jones and Medd, 2000). This approach is reinforced not only by ecological (Davis et al., 2003) but also by economic simulation models (Jones and Medd, 2000). False seedbed technique is a method providing weed seed bank depletion. The principle of flushing out germinable weed seeds before crop sowing forms the basis of the false seedbed technique in which soil cultivation may take place days or weeks before cropping (Johnson and Mullinix, 1995). Germination of weed seeds is stimulated through soil cultivation (Caldwell and Mohler, 2001). Irrigation is suggested to provide the adequate soil moisture required for sufficient weed emergence. In the case of false seedbed, emerged weeds are controlled by shallow tillage operations (Merfield, 2013). Control of weeds and crop establishment should be delayed until the main flush of emergence has passed in order to deplete the seedbank in the surface layer of soil and reduce subsequent weed emergence (Bond and Grundy, 2001).
The knowledge about seed germination for the dominant weed species of a cultivated area is vital for predicting weed seedlings emergence. The possibility of predicting seedling emergence is essential for improving weed management decisions. However, weed emergence is the result of two distinct processes, i.e., germination and pre-emergence growth of shoots and roots, which react differently to environmental factors and should therefore be studied and modeled separately (Colbach et al., 2002a). In temperate regions, soil temperature is probably the most distinct and recognizable factor governing emergence (Forcella et al., 2000). Soil temperature can be used as a predictor of seedling emergence in crop growth models (Angus et al., 1981). Soil temperature can also be used for predicting weed emergence, but only if emergence can be represented by a simple continuous cumulative sigmoidal curve and the upper few centimeters of soil remain continuously moist (Forcella et al., 2000).
The Possible Effects of Light, Gaseous Environment of the Soil, Soil Nitrates Content and Soil PH on Seed Germination of Various Weed Species
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Fluctuating temperatures belong to parameters that can remove the constraints for the seed germination of many weed species once the degree of dormancy is sufficiently low (Benech-Arnold et al., 2000). In particular, the extent and number of diurnal soil temperature fluctuations can be critical in lessening seed dormancy of several species. For example, alternating temperatures at 25 °C increased germination of Amaranthus retroflexus (L.), Amaranthus spinosus (L.), and Amaranthus tuberculatus (L.) from 23 to 65, 8 to 77, and 9 to 57%, respectively, as compared to non-alternating temperatures. Fluctuating temperatures from 2.4 to 15°C can terminate the dormancy situation in Chenopodium album (L.) seeds (Murdoch et al., 1989). Either four diurnal cycles of 12°C amplitude or 12 diurnal cycles of 6°C amplitude were necessary for the emergence of D. sanguinalis (King and Oliver, 1994). The number of cycles of alternating temperatures needed to end the dormancy situation has to be investigated. In Sorghum halepense (L.) Pers., a 50% increase in cycles of alternating temperatures can double the number of seeds that are released from dormancy (Benech-Arnold et al., 1990). Furthermore, if the demand for fluctuating temperatures to terminate dormancy in the seeds of this species is not satisfied, a loss of sensitivity to fluctuating temperatures occurs in a proportion of the population (Benech-Arnold et al., 1988). The variation among weed species regarding the demands for fluctuating temperatures for seed germination points out the need for further investigation regarding the effects of fluctuating temperatures in germination of noxious weed species in different regions around the world and under various soil and climatic conditions.
Germination speed of Alopecurus myosuroides (Huds.) seeds decreased with temperature, whereas the final proportion of germinated seeds was not significantly influenced (Colbach et al., 2002b). Minimum temperature required for seed germination is different for various weed species. Minimum temperature required for seed germination has been estimated at 0°C both for the winter annual A. myosuroides (Colbach et al., 2002a) and the summer annual P. aviculare (Batlla and Benech-Arnold, 2005). However, Masin et al. (2005) estimated the base temperature for Digitaria sanguinalis (L.), Setaria viridis (L.), P. Beauv., Setaria pumila (Poir.), Roem. & Schultes and Eleusine indica (L.), at 8.4, 6.1, 8.3, and 12.6°C, respectively. Moreover, the mean Tb recorded for summer annuals Amaranthus albus (L), Amaranthus palmeri (S. Wats.), D. sanguinalis, Echinochloa crus-galli (L.) Beauv., Portulaca oleracea (L.), and Setaria glauca (L.) was ~40% higher as compared to the corresponding value recorded for winter annuals Hirschfeldia incana (L.) and Sonchus oleraceus (L.). Optimal temperature conditions required for terminating dormancy status vary among different species. For example, Panicum miliaceum (L.) seeds lost dormancy at 8°C while P. aviculare seeds were released from dormancy at 17°C (Batlla and Benech-Arnold, 2005). The two germination response characteristics, Tb and rate, influence a species' germination behavior in the field (Steinmaus et al., 2000). Extended models should be developed to predict the effects of environment and agricultural practices on weed germination, weed emergence, and the dynamics of weed communities in the long term. This requires estimating the baseline temperature for germination for each weed species that are dominant in a cultivated area and recording seed germination in a wide range of temperatures (Gardarin et al., 2010).
Soil moisture is a key parameter affecting the seed dormancy status of many species (Benech-Arnold et al., 2000; Batlla et al., 2004). First of all, the environmental conditions existing during seed development in parent plants and seed maturation affect the relative dormancy of the seeds. Less dormant seeds of Sinapis arvensis (L.) were produced from the mother plants under water stress conditions (Wright et al., 1999) while similar results have been reported regarding either winter annual grass species Avena fatua (L.) or summer perennial S. halepense (Peters, 1982; Benech-Arnold et al., 1992). Moreover, sufficient water potential has been noticed to increase the production of dormant A. myosuroides seeds (Swain et al., 2006).