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Seed Shattering and Dormancy in Weedy Rices

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Seed Shattering and Dormancy in Weedy Rices 45 Chapter 4 Seed shattering and dormancy in weedy rices EARLY AND HEAVY SEED SHATTERING The shattering of seeds as they mature in the inflorescence is the naturally selected trait of plant species that produce dry fruits and seeds and is one of the most important of the seed dispersal and distribution mechanisms in plants. Shattering increases not only the survivability but also the area distribution of the species. Retention of seeds in the inflorescence at least until they all reach maturity is the opposing trait selected at first unconsciously and later consciously by humans in the domestication and improvement of plants to permit and/or facilitate the gathering of the desired plant products: food and feed grains, oilseeds, grain legumes. The selection was at first unconscious as the early human gatherers collected grains retained on the plants for food or seed in preference to those dispersed on the ground. Conscious selection for the non-shattering trait in wheat, barley and rice 6 000–10 000 years ago was probably the first major human “improvement” in plants (Diamond, 1999). Early and heavy seed shattering, the red pericarp and seed dormancy are the constant traits among all the diverse types and other variants of the red rices. If the reference is broadened to include all weedy rices, then seed shattering becomes the invariable trait because some weedy rices, especially in Asia and Africa, have white pericarps, and the seeds of some weedy types are not any more dormant than those of some cultivated varieties. The seeds of Nato, an older-type medium-grain variety, are as dormant as those of many red rice phenotypes for a time after harvest but dormancy is released sooner. Constantin (1960) determined seed shattering for the 1 084 red rice panicles he collected from rice fields in the southwest Louisiana rice area and stated: “No red rice panicles were found that had non-shattering spikelets characteristic of cultivated rice. All had shattering spikelets and no variation among plants was detected.” The older literature on weedy rices in Asia (Graham, 1913; Roy, 1921; Chatterjee, 1947), and red rices in North America (Nelson, 1908; Quereau, 1920; Hodges, 1957; Williams, 1956), and, generally (Grist, 1955), always emphasized the importance of the “shattering”, “shedding” or “deciduous” nature of the spikelets in the persistence and spread of weedy rices. Seed shattering – weedy trait The early and heavy shattering of seeds as they mature in the inflorescence is an important mechanism for their dispersal and distribution. It increases the probability that a substantial portion of the seeds produced by a plant are scattered to the surface of the soil where they can be spread further by wind and water before being consumed by animals, harvested with the grain, or eventually falling to the ground in a clump along with the plant. Shattering is the naturally selected trait of plants but, as mentioned above, it was a very inconvenient trait in plants that produced food grains desired by early humans in the gathering stage of human development. Thus, since the dawn of crop husbandry, non-shattering has been a prized trait in the selection and improvement of varieties of crops. The non-shattering trait has not been disadvantageous to the 46 Weedy rices – origin, biology, ecology and control crops cultivated because their survival is dependent on human activities and not nature. However, the non-shattering trait would be disadvantageous to “wild” plants that grow in association with crops, i.e. weeds. Most of the seeds produced would be removed and destroyed by farmers or mixed in with the harvest and consumed. Farmers plant seeds of crops but not knowingly of weeds. Modern cultural practices and technologies, such as specialized seed production, mechanical harvesting, and grain and seed cleaning, have increased considerably the disadvantages of even relative non- shattering in annual wild plants or weeds growing with crops. Plate 15 compares shattering in panicles from four red rice phenotypes with the non-shattering panicle of the Starbonnet variety. Onset of seed shattering in red rice J.C. DELOUCHE, 1984 While seed shattering is apparently Plate 15 a characteristic of all weedy rices, Top: seed shattering of four RR phenotypes compared with the there is considerable variation in the non-shattering variety Starbonnet (left). Bottom: vigour of RR time and degree of shattering among manifested in tillering as compared with the Starbonnet variety (middle). the phenotypes or ecotypes. Do Lago (1982) determined the onset of shattering for 28 red rice phenotypes collected in Mississippi, the United States of America, in 1980 and found that the mean time of the onset of shattering was 24 days past 50-percent anthesis for all phenotypes, 23 days for the SHR group and 27 days for the BHR group. In 1981, he studied the onset of shattering for 14 selected phenotypes and also determined seed moisture content at the time of shattering (Table 9). The mean shattering time for the BHR types was 3 days later than for the SHR types, but there was much less variation in the onset of shattering (4 days vs 13 days for TABLE 9 Shattering time and sees moisture content of 14 rice phenotypes Type Hull colour Shattering time from 50% anthesis Seed moisture content at start of shattering Mean Range Mean Range (days) (%) Com. var. STW NS NS All reds (13) SH & BH 24 17–30 29.4 23.0–44.2 SHR (9) * STW 24 17–30 27.7 23.0–35.7 BHR (3) BLK 27 25–29 27.6 24.5–31.1 BrHR red (1) * BRN 17 44.2 *Anthesis, shattering and maturation were quite random and irregular in the panicles of two strawhull reds (SHR) and the brownhull red (BrHR). Source: Do Lago, 1982. Chapter 4 – Seed shattering and dormancy in weedy rices 47 the SHR group). Similarly, the mean seed moisture content at the onset of shattering was the same for the SHR and BHR groups but there was less variation among the BHR phenotypes (6.6 percent vs 12.2 percent for the SHR). These data were influenced strongly by two very atypical strawhull phenotypes and the unique brownhull red (BrHR). When they are excluded, the mean seed moisture contents and the ranges at the start of shattering are 26.4 percent, 23–31.1 percent, and 25.8 percent, 23–27.9 percent, for the remaining 10 red rice phenotypes and the 7 SHR types, respectively. In 1981, Do Lago made a more detailed study of the time-course of shattering along the panicle for 6 red rice phenotypes, 4 SHR types and 2 BHR types. Shattering began in the tip of the panicle, as expected, as early as 16 days past 50-percent anthesis for the SHA+ phenotype and as late as 26 days for the 79/11 SHR, but shattering for the latter then progressed to the bottom of the panicle in 2 days (Table 10). The mean times for shattering to begin in the top, middle and bottom third of the panicle for 5 of the 6 phenotypes were 21, 23.4 and 26.4 days, respectively. Seed moisture content of seeds in the upper third of the panicle at the first evidence of shattering ranged from 23.5 to 31.2 percent with a mean of 27.0 percent for the 5 phenotypes. As noted in Table 10, the 79/5 BHR flowered more-or-less randomly rather than from tip to bottom of the panicle so that shattering and maturation were very non-uniform. Thus, the exceptionally high seed moisture content (42.2 percent) at the onset of shattering reflects the mixture of “drier” and “moister” seeds collected in the top third of the plant for the moisture content determination. The 27.0-percent mean moisture for the 5 phenotypes in Table 10 is 2.4 percent lower than that of the 13 phenotypes in Table 9. The difference probably reflects the greater care taken in the detailed study but mostly the inclusion of the brownhull red, coded 78/8, in the mean for the 13 phenotypes. As noted above, it exhibited the same random flowering habit as the 79/5 BHR red excluded from the means in Table 10. Noldin (1995) examined some of the properties associated with seed shattering in 18 red rice ecotypes obtained from four different rice-producing states in the south of the United States of America. Five of the 6 ecotypes from Mississippi had been collected more than 15 years earlier for the studies of Do Lago (1982), Teekachunhatean (1985), Garcia-Quiroga (1987) and others. Noldin found that: ¾ The mean time of the onset of shattering for the 15 ecotypes that exhibited shattering was 15 days after anthesis with a range of 11–18 days. ¾ The mean seed moisture content at the onset of shattering for the 15 ecotypes was 25 percent with a range of 16–30 percent. ¾ The mean shattering index for 17 of the ecotypes on a scale of 1 (very low, < 1 percent) to 5 (moderate, 6–25 percent) to 9 (high, > 50 percent) was 6+ with a range from 1 (3 ecotypes) to 9 (7 ecotypes). TABLE 10 Time-course of seed shattering in selected red rice phenotypes and moisture content of seed in the upper third of the panicle at the first evidence of shattering Phenotype Hull colour Shattering from panicle area Time to reach bottom Seed moisture at top of panicle of panicle Top Middle Bottom (days from 50% anthesis) (days) (%) SHA+ STW 16 18 21 5 31.2 79/1 STW 22 23 25 3 23.5 79/11 STW 26 27 28 2 24.3 79-15 BLK 24 29 33 9 28.3 79/5 BLK 19 20 21 2 42.2¹ 80/1 STW 17 20 25 8 27.9 Mean (less 79/5) 21 23.4 26.4 5.4 27.0 1Anthesis, shattering and maturation were not uniform along the panicle axis.
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