Abstract
Seeds have well-established passive physical and chemical defense mechanisms that protect their food reserves from decay-inducing organisms and herbivores. However, there are few studies evaluating potential biochemical defenses of dormant seeds against pathogens. Caryopsis decay by the pathogenic Fusarium avenaceum strain F.a.1 was relatively rapid in wild oat (Avena fatua L.) isoline “M73,” with >50% decay after 8 days with almost no decay in wheat (Triticum aestivum L.) var. RL4137. Thus, this fungal strain has potential for selective decay of wild oat relative to wheat. To study defense enzyme activities, wild oat and wheat caryopses were incubated with F.a.1 for 2–3 days. Whole caryopses were incubated in assay reagents to measure extrinsic defense enzyme activities. Polyphenol oxidase, exochitinase, and peroxidase were induced in whole caryopses, but oxalate oxidase was reduced, in response to F.a.1 in both species. To evaluate whether defense enzyme activities were released from the caryopsis surface, caryopses were washed with buffer and enzyme activity was measured in the leachate. Significant activities of polyphenol oxidase, exochitinase, and peroxidase, but not oxalate oxidase, were leached from caryopses. Defense enzyme responses were qualitatively similar in the wild oat and wheat genotypes evaluated. Although the absolute enzyme activities were generally greater in whole caryopses than in leachates, the relative degree of induction of polyphenol oxidase, exochitinase, and peroxidase by F.a.1 was greater in caryopsis leachates, indicating that a disproportionate quantity of the induced activity was released into the environment from the caryopsis surface, consistent with their assumed role in defense. It is unlikely that the specific defense enzymes studied here play a key role in the differential susceptibility to decay by F.a.1 in these two genotypes since defense enzyme activities were greater in the more susceptible wild oat, compared to wheat. Results are consistent with the hypotheses that (1) dormant seeds are capable of mounting complex responses to pathogens, (2) a diversity of defense enzymes are involved in responses in multiple plant species, and (3) it is possible to identify fungi capable of selective decay of weed seeds without damaging crop seeds, a concept that may be applicable to weed management in the field. While earlier work on seed defenses demonstrated the presence of passive defenses, this work shows that dormant seeds are also quite responsive and capable of activating and releasing defense enzymes in response to a pathogen.
Highlights
Weeds cause global crop losses through reduced yield, nutrient, and water competition, and by harboring damaging pests and pathogens (Oerke, 2006)
Apparent oxalate oxidase (OxO) activity was reduced by F.a.1 treatment in whole caryopsis assays of wild oat and wheat (Figure 5), the opposite of the other defense enzymes
Lane (2000) reported that, other than the extractable soluble OxO activity in the epidermis, OxO activity in wheat caryopses and seedlings was insoluble, which is consistent with our observations of insoluble activity associated with caryopses (Figure 5). These findings supported our hypothesis that the defense enzymes, polyphenol oxidase (PPO), CHI, and POD would be induced by the seed decay pathogen, F.a.1, in both wild oat and wheat caryopses
Summary
Weeds cause global crop losses through reduced yield, nutrient, and water competition, and by harboring damaging pests and pathogens (Oerke, 2006). Financial losses due to yield decline can be catastrophic, with weeds causing an estimated $20 billion annually in crop damage losses in the U.S (Pimentel et al, 2005). One reason for the persistence of weeds in agricultural systems is the longevity of viable seeds in the soil, known as the soil seed-bank. Wild oat (Avena fatua L.) is one of the 10 worst annual weeds of temperate regions due to its persistence in the soil seed-bank, abundant seed production, and multiple generations (times of emergence) during the growing season (Beckie et al, 2012). Use of herbicides against wild oat has resulted in many cases of herbicide resistance (Owen and Powles, 2009)
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