Abutilon Theophrasti ) Seeds
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Weed Science, 53:479±488. 2005 Corn competition alters the germinability of velvetleaf (Abutilon theophrasti ) seeds Robert E. Nurse Velvetleaf is a troublesome annual weed in many cropping systems of the United Department of Crop and Soil Sciences, Cornell States and Canada. Differences in the growing environment of parent plants can University, Ithaca, NY 14853. Present address: in¯uence the number, structure, germinability, and viability of seeds produced. Thus, Agriculture and Agri-food Canada, Greenhouse and the effects across a range of competitive environments and corn planting dates on Crops Processing Centre, Harrow, ON N0R 1G0, velvetleaf seed production, germination, and seed coat weight were examined under Canada ®eld conditions. Seed production of velvetleaf increased with increasing biomass. Total velvetleaf reproductive output was reduced in competition with corn compared Antonio DiTommaso with monoculture stands. Corn planting date had no effect on the dormancy status Corresponding author. Department of Crop and of seeds, but increased competition from corn resulted in up to a 30% decrease in Soil Sciences, Cornell University, Ithaca, NY 14853; the proportion of seeds that were dormant. Seed and seed coat weights also decreased [email protected] for plants of velvetleaf grown in competition with corn compared with those grown in monoculture. These ®ndings suggest that velvetleaf plants growing in relatively noncompetitive environments, such as along ®eld edges or in ®eld areas with poor crop stands, are likely not only to produce a greater number of seeds but also a greater proportion of seeds that are dormant. This alteration in the dormancy status of velvetleaf seeds in the absence or presence of a crop provides unique opportunities for effective long-term management of the soil seedbank in this species, especially for velvetleaf individuals bordering ®elds or growing in fallow areas that might re- quire more stringent control because of increased seed dormancy. Nomenclature: Velvetleaf, Abutilon theophrasti Medic. ABUTH; corn, Zea mays L. `Dekalb 520RR'. Key words: Germination, competition, maternal environment, photoperiod, seed coat, dormancy. Interspeci®c competition between velvetleaf and corn re- Typically, a high proportion of mature velvetleaf seeds sults in signi®cant economic losses in many North American exhibits physical dormancy, a condition in which a hard seed cropping systems (Spencer 1984). Integrated weed manage- coat prevents imbibition of water necessary for germination ment strategies manipulate interspeci®c competitive inter- (Horowitz and Taylorson 1984, 1985; LaCroix and Stani- actions between crops and weeds to suppress weed growth forth 1964; Warwick and Black 1988). Winter (1960) sug- and reproduction (Swanton and Murphy 1996). For ex- gested that the cutinized palisade layer in velvetleaf seeds ample, Teasdale (1998) showed that increasing corn density was largely responsible for their impermeability to water. to at least 90,000 plants ha21 reduced velvetleaf seed pro- This layer comprises half the thickness of the velvetleaf seed duction by up to 94% compared with a standard planting coat and is discontinuous at the chalazal slit. In members density of 64,000 plants ha21. Moreover, velvetleaf seed pro- of the Malvaceae such as velvetleaf, water entry is only per- duction is often eliminated if velvetleaf emergence occurs mitted through the chalazal slit, which in dormant seeds is after the six-leaf stage of corn under this higher corn pop- blocked by the chalazal plug (Baskin and Baskin 1998). The ulation density because of lower light availability to the opening and closing of the chalazal plug is affected by seed weed. In another study, velvetleaf seed production was re- moisture content (Winter 1960). Horowitz and Taylorson duced from 18,000 to 100 seeds m22 for early- (planted at (1984) showed that the permeability of the seed coat to same time as corn) vs. late-emerging (planted 3 wk after water can be affected by temperature and moisture levels corn) individuals in a corn ®eld in Ohio (Cardina et al. during storage. It is also possible that seed coat imperme- 1995). Similarly, Steinmaus and Norris (2002) reported re- ability can be in¯uenced by environmental conditions ex- ductions in velvetleaf seed output from 44,200 seeds plant21 perienced by parent plants during seed development. Vel- in monoculture stands to 349 seeds plant21 in a corn crop vetleaf plants at a competitive disadvantage relative to corn in California. Interspeci®c competition from soybean [Gly- plants and experiencing reduced light availabilities might di- cine max (L.) Merr.] was also shown to signi®cantly reduce vert resources away from reproductive structures such as the velvetleaf survival and seed production (by up to 82%) rel- seed coat. This could subsequently compromise the integrity ative to velvetleaf plants grown in monoculture (Lindquist of the palisade layer or other components of the seed coat et al. 1995). Although numerous studies have demonstrated and result in seeds that are permeable to water in regions the deleterious effect of crop competition on velvetleaf seed other than the chalazal slit. Although not possessing physical production, no research to date has examined the effects of seed dormancy, buckhorn plantain (Plantago lanceolata L. competitive interactions on the viability and dormancy sta- [Plantaginaceae]) seed coat weight increased relative to em- tus of velvetleaf seeds produced by these plants. bryo/endosperm weight when parent plants were grown un- Nurse and DiTommaso: Velvetleaf seed dormancy x 479 der increasing temperatures (Lacey et al. 1997). Previous No corn was planted for the early (May 8) treatment in work (Nurse, unpublished data) suggested that environmen- 2002 because excessive precipitation prevented the use of tal stresses such as drought and decreased photoperiod dur- planting machinery on the ®eld site. Three velvetleaf seed- ing maturation of velvetleaf seeds might reduce their dor- ling transplant dates were established to provide a range of mancy, although it was unclear whether this was a result of competitive hierarchies between corn and velvetleaf: 14 d increased seed coat permeability or some other mechanism. before corn emergence (DBE), 0 d after corn emergence Late-maturing velvetleaf plants might experience both (DAE), and 14 DAE. The 14 DBE treatment was estab- higher levels of competition from a well-established crop lished by transplanting velvetleaf seedlings (one- to two-leaf and decreasing photoperiod. Flower initiation in velvetleaf stage) into the ®eld immediately after the seeding of the is most rapid under short photoperiods (e.g., 11 h), al- corn. In early May, corn generally takes between 10 and 14 though this species eventually ¯owers under longer photo- d to emerge in central New York state; thus, velvetleaf seed- periods (e.g., 15 h) (Patterson 1995). This shorter interval lings in this treatment had an approximate 14-d growth between maturation, ¯owering, and seed set suggests that advantage over corn. the parent plant will have fewer resources to allocate to re- In early May of each year, the soil was moldboard plowed production. Species whose seeds possess physical dormancy and disked, and the nonselective herbicide glyphosate1 was that germinate early in the season and under longer pho- applied at 0.9 kg ai ha21 before corn emergence to control toperiods might produce a higher proportion of water-im- existing vegetation before planting. At corn planting, fertil- permeable (dormant) seeds than plants grown under shorter izer (10-20-20 N-P-K) was added at 224 kg ha21. Gly- photoperiods. Evenari et al. (1966) found that seeds of phosate-resistant corn cultivar `Dekalb 520RR' was sown for Ononis sicula Guss. (Fabaceae) had thicker, more imper- all corn planting dates at a row spacing of 76 cm and at a meable seed coats when grown under a 20-h photoperiod density to achieve 70,000 plants ha21. compared with seeds produced by plants grown under an 8-h photoperiod. However, few studies have investigated the Seedling Source effect of photoperiod in species whose seeds exhibit physical dormancy (hardseededness). Seeds of velvetleaf used to produce seedlings for this study No studies have examined the combined effects of corn were randomly collected from a corn ®eld at the Robert competition and corn planting date on the viability and Musgrave Agronomy Research facility in September 2001 dormancy of velvetleaf seeds produced under these condi- and were stored dry in the dark at 5 C until required. Only tions. The objectives of this study were to examine the effect seeds of relatively equal weight (9.0±9.5 mg) were used. In of environmental conditions during seed maturation, in- spring 2002 and 2003, seeds were soaked in boiling water cluding competition with corn as well as conditions associ- in cheesecloth for 10 s then placed on ®lter paper moistened ated with different corn planting dates (e.g., photoperiod, with distilled water in petri dishes on a lab countertop at ; temperature) on seed production and the proportion of dor- room temperature ( 22 C). Nearly all seeds germinated mant seeds in velvetleaf. We hypothesized that velvetleaf within 3 d. Individual germinated seeds were placed in peat 2 3 plants grown under stressful conditions (i.e., intense corn pots (40 cm ) containing a 1:1 (v/v) mixture of peat moss competition and short photoperiod) would produce a lower and vermiculite. Pots were transferred to a glasshouse under