A Physiological Disorder of Peppers
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HORTSCIENCE 52(1):4–9. 2017. doi: 10.21273/HORTSCI11123-16 results to interested readers, not only of aca- demic interest but commercial as well. This Capsicum review should facilitate advances toward a A Review of Chile Pepper ( deeper understanding of the disorder leading to effective treatments and additional preven- annuum) Stip: A Physiological Disorder tive measures instead of repeating previously of Peppers conducted, yet difficult to obtain, studies. Description of Lesions James C. Fulton1 Plant and Environmental Sciences, New Mexico State University, Las Symptoms of stip appear as pale to dark Cruces, NM 88003 green (Eijsinga et al., 1973; Hibberd, 1981; Smith et al., 1996, 1997, 1998), yellow Mark E. Uchanski2,3 (Aloni et al., 1994; Feigin and Sagiv, 1987; Horticulture and Landscape Architecture, Colorado State University, Fort Smith et al., 1996, 1997, 1998), or black (Smith et al., 1996, 1997, 1998; Villalon, Collins, CO 80523-1173 1975) lesions in pepper pod tissue. Affected Additional index words. abiotic disorder, environmental stress, bell, chile tissue may be sunken (Eijsinga et al., 1973; Feigin and Sagiv, 1987; Hibberd, 1981; Abstract. Stip, a physiological disorder, which affects certain pepper (Capsicum annuum) Villalon, 1975), oval or irregularly shaped cultivars and pod types, has received limited formal academic attention. The disorder, (Hibberd, 1981; Villalon, 1975). Chlorotic frequently noted for its appearance in bell peppers, but also present in pimento and New or necrotic areas of pod wall cells (Hibberd, Mexican pod types, has been attributed in the literature to nutrient imbalances and/or 1981; Villalon, 1975) have been reported as temperature extremes. Symptoms of this disorder present on fully developed fruits as between 1 and 10 mm in diameter (Eijsinga brown, black, and yellow ovoid subcutaneous chlorotic lesions ’1.3 cm in length and 0.6 cm et al., 1973; Feigin and Sagiv, 1987; Hibberd, in width extending from the endocarp to the mesocarp, without apparent expression 1981; Smith et al., 1996, 1997, 1998; Villalon, in the cuticle. Chile pepper has been significantly affected by stip for the past several 1975), however, observed by the author as decades leading to varying degrees of commercial losses. Due to the sporadic and between 5 and 15 mm in southern New seemingly spontaneous nature of the disorder, prolonged research interest has not been Mexico, that extend throughout the exocarp sustained. This review provides the reader a comprehensive synthesis of all available and mesocarp (Fig. 1). literature regarding this physiological disorder. Especially because research into this Lesions may also have a soft to dry texture disorder is scarce, it is essential that any and all previous research be made available (Villalon, 1975). The severity (number of together in one document to interested scientists, breeders, and growers. To achieve this lesions per symptomatic pod) may vary be- goal, all available stip-related publications were obtained and analyzed; additionally, tween 1 and 30 (Eijsinga et al., 1973; Feigin and searches were performed using all names associated with the disorder. The literature Sagiv, 1987; Hibberd, 1981; Smith et al., 1996, suggests that stip manifestation is a consequence of an interaction between a genetic 1997, 1998; Villalon, 1975). Except for these component (open-pollinated cultivars being more susceptible to symptom expression lesions, the vegetative and reproductive por- than hybrids) and an environmental component (increased nitrogen application, tions of plants appear normal (Villalon, 1975). shading, and soil applications of lime appear to increase both severity and incidence). Description of Symptom Manifestation The terms ‘‘stip’’ (Smith et al., 1996, 1997, 1996, 1997, 1998; Stolk and Maaswinkel, 1977; Stip symptom development has been 1998), ‘‘black spot’’ (Villalon, 1975), ‘‘colour Uffelen, 1973, 1975; Villalon, 1975); however, observed in the greenhouse (Aloni et al., spots’’ (Aloni et al., 1994; Feigin and Sagiv, pimento, and New Mexican pod-type peppers 1994; Carroll, 1979; Eijsinga et al., 1973; 1987), ‘‘green spotting’’ (Carroll, 1979), ‘‘green have also been mentioned (Smith et al., 1995, Stolk and Maaswinkel, 1977) as well as under pitting’’ (Hibberd, 1981), ‘‘spotting’’ (Uffelen, 1996, 1997, 1998; Villalon, 1975; Table 1). field conditions (Feigin and Sagiv, 1987; 1973, 1975), ‘‘bitter pit’’ (Eijsinga et al., 1973), Previous studies have attempted to estab- Hibberd, 1981; Smith et al., 1996, 1997, and ‘‘pepper spot’’ (Stolk and Maaswinkel, lish that stip is not a consequence of any 1998; Villalon, 1975), and its severity and 1977) are different names that have been used known virus, bacterium, or fungus (Eijsinga incidence have been reported as variable from in the literature to describe what is likely the et al., 1973; Hibberd, 1981; Smith et al., year to year and therefore difficult to predict same physiological disorder of peppers. This 1996; Villalon, 1975), and no insect damage (Eijsinga et al., 1973; Smith et al., 1996, disorder, hereafter referred to as stip, was is reported in connection with this disorder 1997, 1998; Villalon, 1975). Although the independently described twice in the 1970s, (Eijsinga et al., 1973; Hibberd, 1981; Villalon, disorder has been reported as most pro- once in the Netherlands and once in Texas 1975). Consequently, symptoms are believed nounced when pods are fully mature and (Eijsinga et al., 1973; Villalon, 1975). Stip to be a manifestation of a physiological dis- red in color (Feigin and Sagiv, 1987; Hibberd, symptoms have been described most exten- order (Aloni et al., 1994; Carroll, 1979; 1981; Smith et al., 1996, 1997, 1998), the sively in bell peppers (Aloni et al., 1994; Eijsinga et al., 1973; Feigin and Sagiv, 1987; disorder has also been noted when pods are Carroll, 1979; Eijsinga et al., 1973; Feigin Hibberd, 1981; Smith et al., 1996, 1997; Stolk mature green, with symptoms becoming and Sagiv, 1987; Hibberd, 1981; Smith et al., and Maaswinkel, 1977; Uffelen, 1973, 1975; more apparent as they mature further (Aloni Villalon, 1975). The disorder has been spo- et al., 1994; Villalon, 1975). Consequently, radically reported in geographically distant symptoms are most often noted during har- Received for publication 5 July 2016. Accepted for academic journals over the past 40 years, vest in the late summer and early fall publication 12 Oct. 2016. likely because it presents as a problem in transition period oftentimes occurring when Funding for this research was provided by the New a given production area for brief, and appar- nights are cool, daylength is shorter, and Mexico Chile Association. Additional salaries and ently random times. As a result, there has not dews are heavy for fall harvested peppers research support were provided by state and federal been sustained and focused attention on study- (Aloni et al., 1994; Eijsinga et al., 1973; funds appropriated to the New Mexico Agricultural ing the condition. Therefore, a comprehensive Experiment Station. Feigin and Sagiv, 1987; Hibberd, 1981; 1Graduate Research Assistant. review that consolidates the information gen- Smith et al., 1996, 1997, 1998; Villalon, 2Assistant Professor. erated through these spatially and temporally 1975; Wien and Zitter, 2003). Increased 3Corresponding author. E-mail: mark.uchanski@ disparate studies can be useful to describe, symptom severity and incidence have been colostate.edu. synthesize, and present previous research described during the middle portion of 4 HORTSCIENCE VOL. 52(1) JANUARY 2017 REVIEW Table 1. A summary of the published literature referencing the physiological disorder known as stip in peppers, arranged alphabetically by author. Author (yr) Location Pod type, cultivar Environment Fruit stage Analysis type Aloni et al. (1994) Israel Bell; Maor, Lady Bell, and 899 Greenhouse Mature red V (N, K, Ca, photosynthesis); (increasing in resistance) R (N, K, Ca, Mg, oxalic acid, yield, severity) Carroll (1979) Australia Bell; California Wonder Greenhouse NA R (incidence, Ca, B, yield) Eijsinga et al. (1973) Netherlands Bell Greenhouse Mature red R (Na, K, Ca, Mg, P, Cl, N, NO3-N, S, SO4-S, Mn, Fe, Al, B); soil components affecting manifestation Feigin and Israel Bell; Maor Field Mature red V (N); R (N, incidence/severity) Sagiv (1987) Hibberd (1981) Australia Bell; Sheba, Yolo Y, Midway, Field Mature red R (yield, incidence/severity) Early Bountiful, Allbig, Green Giant, Super Set, Jade, Yolo Wonder, California Wonder, Miss Belle Mullen et al. (2000) California Bell; Emerald Isle, Grande Rio, Field NA R (incidence, yield) Bigga-Pep, Early Sunsation, HA-65, P33F1 Smith et al. (1995) California Bell; Galaxy, King Arthur (R), Field Mature red V (N, K, Ca); R (N, K, Ca, Yolo Wonder L, Grande Rio (S) incidence/severity) Smith et al., (1996) California Bell; Galaxy, King Arthur, Field Both V (N, K, Ca, Mg, Na, B); Grande Rio 66, Jupiter (S) R (N, K, Ca, Mg, Na, B, incidence/severity) Smith et al. (1997) California Bell; Grande Rio 66 (S) Field Mature red V (NO3-N, N, P, K, Ca, Mg, Na, Cl, B, Zn); R (N, P. K, Ca, Mg, Na, Cl, B, Zn) Smith et al. (1998) California Bell; Yolo Wonder A, Yolo Wonder B, Field Mature red V (N, K, Ca, Mg, Na); Jupiter, Keystone Resistant Giant, R (incidence/severity, yield) Grande Rio 66, Mercury, Pimlico, Loribelle, Cal Wonder 300 (R), Capistrano, Merced, Emerald Giant, Taurus, Gusto, D-13, Pip Smith et al. (1999) California Bell; Cal Wonder, Gusto, Greenhouse and Field Mature red R (incidence/severity) King Arthur, and Grande Rio Stolk and Netherlands Bell; Propa, Deltana, No. 40, Manto, Greenhouse Mature red R (yield, severity) Maaswinkel, 1977 Rumba, Bell Boy Uffelen (1973) Netherlands Bell Field Mature red R (Na, K, Ca, Mg, P, Cl, N, NO3-N, S, SO4-S, Mn, Fe, Al, B); soil components affecting manifestation Uffelen (1975) Netherlands Bell; No. 938, Bell Boy, Field Mature red R (incidence/severity, yield) Rumba, Deltana, Improved Glass, Sonette Villalon (1975) Texas Bell. Also mentions Anaheim Plants transplanted from Both Plant inoculation and (New Mexico pod type) field to greenhouse pathogen identification NA = not available.