ment with the predictions we devel- oped in 1993. One of the most encour- aging observations is that three of the populations exhibiting the greatest re- sistance to the SLWF were also high yielding, based on the first year of yield data. We will continue to select for SLWF resistance and improved forage yield. We are hopeful that we will also see a significant increase in yield once we develop populations that exhibit SLWF resistance levels below the as- sumed economic-threshold level of 2.0. Conclusion Cultural management of the SLWF in alfalfa by either chemical control or cutting management is not feasible. The number of days is susceptible to depends on the planting date. We have developed plant-breeding methodology to successfully select for genetic resistance to the SLWF. Screen- ing is conducted under field condi- Imperial Valley conditions tions in the Imperial Valley during August and September. is pro- duced on the selected plants between limit Karnal bunt in wheat September and March in a ”winter” nursery in Chile. This permits two Gerald J. Holmes 0 Lee F. Jackson o Thomas M. Perring generations per year and complete pest and yield evaluation at more than The amount of disease occurring Karnal Bunt (KB) is a minor disease one location starting in the spring after in any given area depends on the of wheat that until March 8,1996, the year of selection. Taking into ac- presence of the pathogen in suffi- was of little concern to the US grain count both our early predictions and cient abundance, susceptible industry. On that date, scientists at our current progress, we expect that hosts and favorable climatic con- USDA’s Agricultural Research Ser- populations with economic resistance ditions. Each of these factors vice confirmed the presence of to the SLWF will be selected this fall. were severely limiting to the es- Karnal bunt in Arizona-certified du- We are working to provide California tablishment of Karnal bunt (KB) in rum wheat seed (cv. Reva), produced growers with resistant cultivars as the Imperial Valley during the near Gila Bend, Ariz. Seed from in- rapidly as possible. 1996 growing season, and none of fested seed lots was planted in Ari- the 1,476 fields (106,592 acres) zona, New Mexico, Texas and Cali- L.R. Teuber is Professor and K.L. Taggard tested was shown to be infected fornia. Fourteen fields amounting to is Staff Research Associate, Department of with KB. Karnal bunt does not ap- about 960 acres, primarily of the cul- Agronomy and Range Science, UC Davis; tivar Kronos, were planted with in- pear to be a threat to wheat pro- L.K. Gibbs is Staff Research Associate, De- fected seed in the Imperial Valley of duction in the Imperial Valley be- partment of Agronomy and Range Sci- California. As part of a larger effort cause desert conditions are ence, Desert Research & Extension Cen- to prevent the spread of KB, wheat in ter; and M.E. Rupert is former Graduate unfavorable for its development. the Imperial Valley was placed un- Research Assistant, Department of However, this does not rule out der a federal quarantine. Fields in Agronomy and Range Science, now Staff the possibility that low levels of the quarantine area had to be tested Research Associate, Department of Pomol- disease may occur occasionally. for KB prior to harvest and again be- ogy, UC Davis. Assuming that current growing fore shipment. All preharvest The authors wish to express their sin- conditions continue and that KB- samples taken from fields in the Im- cere appreciation to the Imperial Whitefly free seed is planted, the Imperial perial Valley (1,476 fields) tested Management Committee for funding the Valley is at low risk for a KB negative for the presence of KB research reported in this paper. outbreak. .

CALIFORNIA AGRICULTURE, MAY-JUNE 1996 29 Distribution and severity Diseased ear-head Karnal bunt is indigenous to India, first being reported near the town of Karnal in 1931. The disease also occurs Secondary sporidia in Syria, Lebanon, Iraq, Pakistan, Af- ghanistan and Mexico. It affects com- mon wheat and to a lesser degree du- rum wheat, and . Estimates @Flowering stage I' ../ a; of loss due to KB are difficult since the disease can reduce seed quality but usually only minimally affects grain w yield. Since the disease is seedborne I1 1) Infected kernels I( I( and many countries have a zero toler- ance policy, indirect costs are incurred I in preventing its spread, while losses I in marketing opportunities arise be- cause of its presence. Karnal bunt can be a severe disease, and losses ranging from 5 to 20% have been reported. However, even during years of severe epidemics in India, the total damage to the wheat crop ranged from 0.3 to 0.5% of total production. Symptoms Karnal bunt usually is noticed first when broken and/or partially smutted kernels are seen in threshed grain. Fig. 1. Life cycle of Karnal bunt infection (Adapted from Joshi et al. 1983). However, symptoms are present at the soft-dough stage in the form of black- ened areas surrounding the base of the grain, which extend upward along the suture to varying degrees. Unless the disease is severe, only a few florets per spike are affected and diseased spikes are not conspicuous because the glumes are not distorted by infected kernels. In severely infected spikes, however, the glumes may spread apart near maturity, exposing the bunted grains. Infected grains are irregularly distributed in the spike; some are com- pletely infected, but most are partially infected (hence the alternate common name of the disease, partial bunt). The A This wheat spike shows glume spread caused by Karnal bunt infection. In se- weight of infected grains is directly re- verely infected spikes, the glumes may lated to the severity of infection; as the spread apart near maturity, exposing degree of infection increases, the bunted grains. weight of the grains correspondingly decreases. While diseased often retain a 4 Bunted wheat kernels infected with Karnal bunt are usually first noticed in partial seed coat, the embryo and part 01 7 threshed grain. of the can be converted to

30 CALIFORNIA AGRICULTURE, VOLUME 51, NUMBER 3 masses of small black , which emit a fishy odor (due to the presence of trimethylamine) similar to common bunt. Partially bunted seeds often re- tain their capacity to germinate, and grains with trace-to-moderately-low infections may produce healthy plants. The causal organism indica Mitra (syn. Neovossia indica [Mitra] Mundk.) is in the Class Basidiomycetes, Order Ustilaginales, Family Tilletiaceae. Teliospores are 22 to 49 pm in diameter, more or less spherical, black in color and have a slightly rough surface. The teliospores are nearly twice the size of those of the related species and Tilletia confroversa, causal organisms of common and dwarf bunt, respectively. The teliospores of KB bear truncate projections surrounded by a delicate membranous sheath. Germinating teliospores form promycelia and produce numerous haploid, sickle-shaped sporidia (fre- quently more than 100) in a whorl at the apex. The sporidia do not fuse but germinate directly to form hyphae and/or uninucleate alantoid second- ary sporidia, which are forcibly dis- charged into the air. Primary sporidia have dimensions ranging from 64 to 79 pm in length and 1.6 to 1.8 pm in width. Secondary sporidia have di- mensions ranging from 12 to 13 pm in length and about 2 pm in width. Disease cycle Sources of inoculum are diseased seed and soil contaminated with spores from previous crops (fig. 1). The delicate pericarp surrounding each sorus of teliospores is easily bro- ken during harvest, dispersing the spores to contaminate the soil. There are no data available to determine the relationship between the load per seed and subsequent disease appearance in the field. Environment plays a primary role in disease devel- opment (Smilanick et al., 1985). The teliospores germinate in response to

CALIFORNIA AGRICULTURE, MAYJUNE 1996 31 free moisture and produce numerous though a low level (6.7%)of germina- In the Imperial Valley, wheat is sporidia at the soil surface that are tion of teliospores from freshly har- planted between Nov. 15 and March 1. forcibly ejected and then dispersed by vested kernels has been observed after Most is planted between late Novem- wind, splashing water and insects. 1 week of storage. Teliospores remain ber and Jan. 1, and in 1996,71%was Sporidia have a short life-span, viable in the soil for up to 45 months. planted before Jan. 1. Due to the heat even at high relative humidity They survive longer when buried in unit differences between winter and (Smilanick et al., 19891, and generally dry soil than in moist, cropped soils or spring, the earlier wheat is planted, survive for only a few hours when air- on the soil surface. the longer its growing season. For ex- borne. Long-distance dispersal of sec- ample, wheat planted on Nov. 15 re- ondary sporidia probably can occur Susceptible period quired 5 more days to reach heading only during moist periods and under In the Imperial Valley, the fields stage than wheat planted on Dec. 15. cloud cover at night. Plants are most that are most vulnerable to KB are Similarly, wheat planted on Jan. 13 re- susceptible to infection when spikes those that were previously contami- quired 17 fewer days to reach heading emerge from the boot, but infection nated with KB spores, were planted than a Dec. 15 planting. Combining can occur throughout the flowering with infested seed or are planted to these data with performance data on period. Sporidia infect the ovaries, di- (i.e., hard red wheat, the three most popular wheat rectly penetrating the glumes and which is more susceptible to KB than varieties grown in the valley, we de- ovary wall. Infection can move across durum wheat). In 1996,14 fields (960 termined the time to heading as a all florets of a spikelet and to the acres) were planted with infested function of planting date (table 1). spikelets above and below the initial wheat seed, and all tested negative for Days to heading is the number of infection site. Diseased kernels may be KB teliospores. In the same year, less days from planting until 50% of the partially or completely displaced by than 1,000 acres of common wheat spikes are fully emerged (peduncles masses of teliospores in cool, humid or were planted in the valley; all of these visible). The period from the begin- wet weather. Sporulation starts at the fields also tested negative. Of the re- ning of spike emergence (the begin- embryo end and moves along the su- maining wheat acreage, approximately ning of the KB-susceptible period) to ture of the grain. The extent of kernel 90% was planted with one of 3 durum the completion of spike emergence bunting depends on when infection varieties: Kronos (60%), Westbred Turbo (heading date) is temperature-depen- occurs and how long favorable condi- (20%), and Yavuros 79 (10%). dent and is approximately 5 days for tions last. Unlike other and bunt fungi, the Jan. 13 planting, 10 days for the Fresh teliospores require a dor- T. indicu does not infect seedlings. This Dec. 15 planting, and 14 days for the mancy period of up to 6 months, al- must infect plants between the Nov. 15 planting. The number of days beginning of spike emergence and the between the heading date and the completion of flowering or anthesis (the end of the KB-susceptible period) is influenced less by temperature, and is estimated to be 5,6 and 7 days for planting dates of Jan. 13, Dec. 15 and Nov. 15, respectively. Utilizing infor- mation in Table 1, along with the above adjustments for days to the be- ginning of spike emergence and to the end of flowering (period of KB-suscep- tibility), we estimated the KB-suscep- tible period to be 10 days, 16 days and 21 days for planting dates of Jan. 13, Dec. 15 and Nov. 15, respectively. For the three durum varieties, which vary in their time to heading, the KB-susceptible period was calcu- lated for three planting dates (table 2). Taking these as representing the ma- jority of wheat varieties planted, and the three planting dates as representa- tive of the Imperial Valley planting

32 CALIFORNIA AGRICULTURE, VOLUME 51, NUMBER 3 times, the KB-susceptible period in Considering the conditions that must 1996 was estimated to begin Jan. 8 and be met for KB disease development, it end March 22. To be conservative and is not surprising that KB is a minor accommodate the few earlier wheat disease even in areas where it is en- plantings, we considered the period of demic. While it is not impossible for KB Jan. 1,1996, through March 31,1996, to occur in the Imperial Valley, the prob- as the KB-susceptible period. ability of its occurrence is very low.

Weather (Jan. 1-March 31,1996) G.J. Holmes is former Imperial County - Dr. D.V. Singh of the Indian Agri- .$ Cooperative Extension Pathology Farm n culture Research Institute has collected g Advisor, UCCE Imperial County, and meteorological data in India during currently Professor and Extension Plant outbreaks of Karnal bunt. These data Pathologist, Department of Plant Pathol- Karnal bunt is associated with a maximum ogy, North Carolina State University, Ra- associate the occurrence of KB with temperature of 74°F and relative humidity the following weather parameters higher than 80%. In 1996, these conditions leigh, N.C.; L.F. Jackson is Extension (Royer, 1984): rarely persisted longer than 24 hours in Agronomist, Department of Agronomy the Imperial Valley. and Range Science, UC Davis; and T.M. weekly mean maximum tempera Perring is Professor, Department of Ento- tures: 67"-74°F (19"-23"C) and humidities. Daily temperatures on mology and Coordinator for Planning, Insti- weekly mean minimum tempera Jan. 28 and 30, Feb. 1,2,3, 10, 11,12 tute for Desert Agriculture, UC Riverside. tures: 46"-54°F (8"-12"C) and 21, and March 4,5,12 and 13 fell weekly mean relative humidity: within the temperature range associ- 80% References: ated with KB reported by Singh et al. Another report (Singh et al. 1983) (1983). Of these dates, maximum hu- Aujla SS, Sharma K, Chand K, Sawney states, "If at the time of anthesis, the midities were above 70% on Jan. 28 SS. 1977. Influence of weather factors temperature ranges between 67"-74°F and 30, Feb. 1,2,3,10,11 and 12, and on the incidence and epidemiology of and 46"-50°F (19"-23"C and S0-lOoC) March 13. Minimum relative humidi- Karnal bunt of wheat. Indian J. Ecol. 8: coupled with a slight shower provid- ties on these days always dipped be- 175-9. ing high atmospheric humidity, the low 70%, the minimum humidity asso- Joshi LM, Singh DV, Srivastava KD. disease gets established very well." In ciated with KB reported by Aujla et al. 1983. Karnal bunt: A minor disease the Imperial Valley, a total of 3 mm of (1977). Rain fell in the Imperial Valley that is now a threat to wheat. The Ba- precipitation fell between Nov. 1 and only on Feb. 12 (1 mm) and March 1 tanical Review 49:309-30. May 6 (1 mm in mid-February and 2 (2 mm). Even considering the microcli- mm in early March). Other studies matic conditions at the soil surface cre- Royer M. 1984. Travel to India to re- note "the high incidence of the disease ated by irrigated cropping systems, we view Karnal bunt research. USDA, ... when the temperature ranges from conclude that conditions required for North Atlantic Area Plant Disease Re- 64"-73°F (18"-22.5"C) and relative hu- KB to occur rarely persisted longer search Lab. May 30,1984. midity is above 70%" (Aujla et al. than 24 hours in the Imperial Valley in Singh DV, Joshi LM, Srivastava KD. 1977). While these weather data are 1996. 1983. Karnal bunt - A new threat to very general, they provide some mea- wheat in India. Recent Advances in Summary sure for comparison with the 1996 Im- Plant Pathology. Print House (India) perial Valley weather conditions. The amount of disease occurring in Lucknow: 121-35. The KB-susceptible period was or on any host is dependent upon the shortened by rising temperatures in presence of the pathogen in sufficient Smilanick JL, Hoffmann JA, Royer the Imperial Valley beginning around abundance, a favorable environment MH. 1985. Effect of temperature, pH, March 1. During this period, tempera- and a susceptible host. For Karnal light, and desiccation on teliospore germination of Phytopa- tures were in the upper 70s to low 80s bunt of wheat in the Imperial Valley, Tilletia indica. (higher than Singh's maximum range) all three factors were at a minimum in thology 75: 1428-31. and maximum relative humidity was 1996. The amount of pathogen intro- Smilanick JL, Prescott JM, Hoffmann between 65 and 75% (lower than duced was limited to a small number JA, Secrest LR, Wiese K. 1989. Envi- Singh's reported humidity require- of spores on wheat used to plant 960 ronmental effects on survival and ment). Perhaps more important than acres, and the weather conditions, growth of secondary sporidia and maximum and minimum tempera- planting dates and varieties were not teliospores of Tilletia indica. Crop Pro- tures was the duration of temperatures conducive to teliospore germination. tection 8: 86-90.

CALIFORNIA AGRICULTURE, MAY-JUNE 1996 33