Observations of pollen tube growth in Nicotiana alata and their implications for the mechanism of self-incompatibility

 Style squashes and stylar grafts were used to examine the growth of Nicotiana alata pollen tubes in self-compatible and self-incompatible styles. Compatible tubes typically showed a uniform layer of callose deposition in the walls and in small plugs spaced at regular intervals within the tube. Incompatible tubes were characterised by the variability of callose deposition in the walls and by larger, closer and more irregularly spaced plugs. There was no difference in the growth rate of compatible and incompatible tubes during growth through the stigma, but within the style most compatible tubes grew 20–25 mm day^-1 (maximum 30 mm day^–1), whereas incompatible tubes grew 1.0–1.5 mm day^-1 (maximum 5 mm day^–1). Many incompatible tubes continued to grow until flowers senesced, and only a small proportion died as a consequence of tip bursting. Grafting compatibly pollinated styles onto incompatible styles showed that the incompatible reaction could occur in pollen tubes between 2 and 50 mm long, and that inhibition of pollen tube growth occurred in both the upper and lower parts of the transmitting tract. Grafting incompatibly pollinated styles onto compatible styles showed that the incompatible reaction was fully reversible in at least a proportion of the pollen tubes. The findings are not consistent with the cytotoxic model of inhibition of self-pollen tubes in solanaceous plants, which assumes that the incompatible response results from the degradation of a finite amount of rRNA present in the pollen tube. However, if pollen tubes do in fact synthesise rRNA, the findings become consistent with this model. Received: 23 May 1996 / Revision accepted: 22 August 1996     

Analysis of stylar self-incompatibility competence by use of heat induced inactivation

Summary Immersion of Lilium longiflorum pistils in 49 °C water for increasing durations of 1,2,3, or 4 minutes immediately prior to incompatible pollination resulted in a correspondingly progressive decrease in the stylar self-incompatibility competence, as determined from the lengths attained by pollen tubes during 48 hours growth in the styles at 24 °C. Neither pistils remaining on the plant nor those detached from the plant which were immersed after anthesis in 49 °C water for 5 minutes regained self-incompatibility competence during a 48 hour incubation at 24 °C prior to incompatible pollination. Heat treatment of detached pistils as early as 39 hours prior to bud anthesis also resulted in an inactivation of stylar self-incompatibility competence when incompatible pollination was made at 24 hours after anthesis. Experiments utilizing heat treatment of partial lengths of detached whole styles revealed that pollen tubes which have grown through as much at 45 millimeters of either a physiologically incompatible or compatible portion of the style are still capable of shifting to either a higher growth rate or lower growth rate upon entry into respectively either a physiologically compatible or incompatible portion of the style.     

Post‐pollination process in a partially self‐compatible distylous plant,Primula sieboldii (Primulaceae)

Pollen germination and pollen‐tube growth under natural conditions were observed in a population of a distylous species, Primula sieboldii, in which partial self‐compatibility has been demonstrated in some long‐styled genets. We observed post‐pollination processes microscopically in styles collected after self‐morph and inter‐morph hand pollination (with standardized pollen load on the stigmas) in four genets each from the following three ‘genet types’: self‐incompatible long‐styled (SI), partially self‐compatible long‐styled (SC) and self‐incompatible short‐styled morph genets. Irrespective of the genet type, pollen germination began within 24 h after pollination and tubes of pollen reached to the style base with 48–96 h after inter‐morph pollination. Although pollen tubes germinated after self‐pollination in the SC genets, the number of germinated pollen tubes was significantly lower than in the case of inter‐morph pollination. Few pollen tubes germinated after self‐pollination of the SI or short‐styled genets. In SC genets, the rate of pollen‐tube growth did not differ between self‐morph and inter‐morph pollination (～1.9 mm/day). Therefore, differences in self‐compatibility between SC and SI genets in P. sieboldii are likely to be attributable to differential pollen germination rates rather than to differential pollen‐tube growth rates.     

梨远缘花粉原位萌发及生长特性

应用荧光标记方法对梨远缘花粉在‘丰水’和‘噢嗄二十世纪’柱头上萌发及花粉管生长特性进行观察,结果表明:(1)梨远缘花粉均能在柱头上萌发,但其萌发率不同,授粉后24 h,在‘丰水’柱头上‘红叶桃’花粉萌发率最高,达62.8%,而‘盖县大李’花粉萌发率仅为12.0%,各种远缘花粉在‘丰水’柱头萌发率均高于‘噢嗄二十世纪’柱头。(2)各种远缘花粉管在梨柱头或花柱内生长情况也有差异,‘红叶桃’等核果类花粉管在梨柱头上均表现为扭曲、盘绕等现象,不能穿过柱头;‘红星’和‘红富士’花粉管虽然有少量穿过柱头,但不能进一步在花柱内生长,表现为扭曲变形、先端膨大等不亲和性现象。因此,梨与远缘果树杂交不亲和在柱头上就已发生,这与梨自交不亲和反应发生在花柱内的现象不同。     

GROWTH OF POLLEN TUBES OF OENOTHERA ORGANENSIS THROUGH OTHERWISE INCOMPATIBLE STYLES

Hecht , A. (Washington State U., Pullman.) Growth of pollen tubes of Oenothera organensis through otherwise incompatible styles. Amer. Jour. Bot. 47(1) : 32—36. Illus. 1960.–It has previously been reported that pollen tubes of Oenothera organensis fail to grow from compatible stylar tissue into a graft of stylar tissue containing the same “S” alleles as are present in the pollen tubes. Upon this basis it has been assumed that the incompatibility reaction in this species is equally effective at all levels of the stigma and style. Results contrary to these were obtained by employing a new culturing and grafting technique: styles with attached stigmas were excised from their ovaries and cultured in large Petri dishes. The styles were cut usually 15 mm. below the stigmas, then grafted to similarly cut styles from a plant of another incompatibility class and held together by means of a “splint” made of lactose-gelatin and a square of moistened lens paper. This “splint” was generally effective in keeping the cut ends in contact and in allowing pollen tubes to grow through the scion portion into the stock. In the 95 grafts involving a scion having no “S” alleles in common with those of the pollen and a stock containing the “S” allele of the pollen, tubes grew an average of 22.1 mm. into the stock during a 15-hr. period at 27°C., with some growing as far as 55 mm. under these conditions. These results suggest the possibility of 3 alternative mechanisms: (1) the incompatibility reaction is stronger in the stigma than in the style; (2) in the growth of pollen tubes through compatible tissue some stimulus is received or some substance is formed. which allows continued growth into the otherwise incompatible tissue of the stock; or (3) some of the stylar, inhibitory substance is lost by diffusion into the gelatin mixture at the graft juncture.     

The influence of RNA-synthesis inhibitors on in vivo pollen tube growth and the self-incompatibility reaction in Lilium longiflorum thunb

Summary Injection of 3 X 10^–4 g/ml acridine orange, 2 X 10^–5 g/ml actinomycin-D, or 1.34 X 10^–3 g/ml 6-methyl-purine in stigmatic exudate into the hollow styles of Lilium longiflorum immediately before, or 6 or 12 hr after pollination restricted compatible pollen tube growth to lengths typical of incompatible tubes. Treatment 24 hr after pollination resulted either in no effect on compatible tube growth or in a significant retardation of compatible tube growth but not to the length of tubes in incompatibly pollinated styles. Aqueous solutions of these inhibitors at the same concentrations injected into lily styles 6 or 12hr before pollination significantly stimulated incompatible pollen tube growth. These results suggest that RNA synthesis in the style is necessary for the self-incompatibility reaction and RNA synthesis in the pollen tube is required for compatible tube growth. The RNA synthesis in the pollen tube may be completed by 24 hr after pollination.Paper number 7439 of the Scientific Journal Series, Minn. Agr. Exp. Sta., St. Paul, Minn. The author gratefully acknowledges the support of the Graduate School, Univ. of Minn. and the United Bulb Co., Mount Clemens, Mich.     

Temperature effects on self incompatibility in Lilium longiflorum

Summary Detached pistils of the clonal variety, Lilium longiflorum Aral No. 5, were submerged before pollination in 50°C water for 0, 1, 2, 3, 4, 5, 6 or 7 min and then immediately compatibly and incompatibly pollinated. Incompatibility, as indicated by pollen tube length after 48 h at 23.5°C, was eliminated by a 1–2 min submersion while compatibility was removed by a 4–5 min one. The window of incubation temperatures at which incompatible and compatible pollen tubes are clearly differentiated occurred between 15 and 30°C.     

Pollen tube growth in pistils of female-sterile and fertile plants ofOenothera mut.brevistylis

Summary Pollination and pollen tube growth was observed in pistils of fertile and female-sterile plants ofOenothera mut.brevistylis. The stigmas and styles of sterile plants were very irregularly formed and additional ovules often developed in the style. The malformations did not inhibit pollen tube growth but made it slower and less efficient. The pollen tubes grew in the ovary but did not find the ovules. Near their tips, pollen tubes branched or meandered without any clear direction. Lack of tropism caused by malfunction of the ovules is postulated in the female-sterile plant.     

Studies on heteromorphic self-incompatibility systems: Physiological aspects of the incompatibility system of Primula obconica

Summary In Primula obconica, a species with a heteromorphic self-incompatibility system, the distinction between compatible and incompatible pollen tubes takes place on the stigma surface in thrum flowers, self tubes growing randomly over the papillar cells. No differences were seen between self and cross tube behaviour on the pin stigma surface, but self tubes were inhibited within the stigmatic tissue with differences in tube length evident after 24 h. The stigma surface bears a proteinaceous pellicle and binds the lectin Concanavalin A. Removal of the stigma removes the incompatibility barrier in mature gynoecia. Bud pollination shows that pollen tubes cannot grow in a normal manner on immature stigmas; the random growth of tubes over the stigma surface resembles that of mature thrum selfs. Fewer compatible tubes reach the style base of young gynoecia and smaller numbers of seeds are set than in mature flowers. Pin and thrum pollen grains germinate and grow in aqueous media, thrum tubes growing longer than pin. The presence of H₃BO₄ and CaCl₂ in the growth medium promotes tube elongation and lengths equivalent to compatible styles can be obtained. The pollen grains have proteinaceous materials in their walls which diffuse out on moistening. Prolonged washing in aqueous media removes these materials but the incompatibility reaction remains unchanged. Thus the incompatibility reaction is between pollen tubes and stigmatic tissue and differs from the homomorphic, sporophytic system where pollen wall proteins elicit the incompatibility response.     

Direct pollination of Zea mays ovules in vitro with Z. mays,Z. mexicana and Sorghum bicolor pollen

Summary Sorghum (Sorghum bicolor) pollen tubes penetrated and grew in corn (Zea mays) styles. The limited length of the sorghum pollen tubes (3–5 mm) and the absence of stigmatic hairs on the basal (5–10 mm) section of the corn styles prevented effective pollination in vivo and in vitro. Normal fertilisation occurred after in vitro pollination of exposed corn ovules with either corn or teosinte (Zea mexicana) pollen. Six per cent of corn ovules pollinated directly with sorghum pollen responded by rapid, massive growth of nucellar tissue.     

INCOMPATIBILITY AND THE POLLEN ECONOMY OF JEPSONIA PARRYI

Jepsonia parryi (Saxifragaceae) has heterostylous flowers and is strongly self-incompatible. Pin flowers have long styles, large stigmas, short stamens, and numerous, small pollen grains with finely sculptured walls. Thrum flowers have short styles, small stigmas, long stamens, and fewer, larger pollen grains with coarsely sculptured walls. Pin plants and thrum plants occur in a 1:1 ratio in field populations. Although the insect pollinators of J. parryi transfer ample compatible pollen to pin and thrum stigmas to account for full seed production, much of the pollen deposited on stigmas is incompatible. Analysis of the pollen deposits on stigmas collected from field populations indicates that compatible “legitimate” pollination of pin and thrum flowers is essentially random and is not obviously aided by floral dimorphism. It is suggested that although heterostyly had a positive adaptive value in the past evolutionary history of Jepsonia it is no longer adaptive under the present pollination regime, although it is maintained because of its strong genetic fixity.     

Effects of prior self-pollination on outcrossed seed set in tristylous Pontederia sagittata (Pontederiaceae)

The potential inhibitory effects of incompatible pollen on outcrossed seed set were investigated in mass-flowering, self-incompatible, tristylous Pontederia sagittata. Prior application of self pollen, followed after 2, 4, or 6 h by compatible pollen, was conducted on five genotypes of each of the three style morphs under uniform glasshouse conditions. The greatest reductions in seed set occurred in pollinations of the long-styled (L) morph at the 6 h time interval. Smaller reductions were also found for this treatment in the mid-styled (M) morph. No significant reductions in seed set were observed in the short-styled (S) morph or in the other morphs at shorter time intervals. Observations of pollen germination and pollen tube growth indicated that the lack of inhibitory effects in the S morph may occur because relatively few pollen grains adhered to stigmas in selfpollinations. In the L and M morphs, early germination of self pollen may cause physical clogging of the stigma and style, resulting in a reduced number of compatible pollen tubes in styles. Observations of the structural integrity of styles indicated that prior germination of self pollen resulted in more rapid onset of pistil senescence, particularly in the L morph. These influences may contribute to the morph-specific differences in seed set observed following prior self-pollination of outcrossed flowers. The negative effects of incompatible pollen are likely to be most evident where ecological factors cause delays in the delivery of outcross pollen to stigmas.     

Growth of Trifolium pratense L. Pollen tubes in compatible and incompatible styles of excised pistils

Summary The excised pistil technique was used to study effects of pollen treatments on self-incompatibility in red clover. Pollen was treated directly with infrared or ultraviolet radiation, and indirectly with chemicals that were applied to the stigmas before pollination. The chemicals (boric acid, calcium nitrate, colchicine, cyclohexamide, gibberellic acid, glycine, and indole butyric acid) were each used at five concentrations. High dosage rates of most treatments inhibited pollen growth in styles that were genetically compatible, but none of the treatments had a specific effect on the self-incompatibility mechanism.The effects of growth of compatible or incompatible pollen tubes in a style on subsequent growth of both types of pollen tubes in the same style were studied in an experiment with 6 hr between pollinations. The passage of some compatible pollen tubes through the style did not influence either self- or cross-compatibility in the style after the second pollination. The passage of some self-incompatible tubes into the styles caused a slightly higher number of incompatible tubes to penetrate the style after the second pollination. Each pollen tube appeared to traverse the style independently of other tubes.Cooperative research by Crops Research Division, Agricultural Research Service, U.S. Department of Agriculture, and the Kentucky Agricultural Experiment Station. This article (70-3-163) is published with approval of the Director of the Kentucky Agr. Exp. Sta.     

EFFECT OF FLORAL AGING ON THE GROWTH OF COMPATIBLE AND INCOMPATIBLE POLLEN TUBES IN LILIUM LONGIFLORUM

Compatible and incompatible pollen tube growth in detached styles of three Lilium longiflorum cultivare varies with the physiological age of the style. Before anthesis both compatible and incompatible pollen tubes grow, in 48 hr, only a fraction of the distance compatible tubes grow after anthesis. Incompatible pollen tubes are restricted to about half the distance of compatible tubes in the four days postanthesis, but thereafter increase up to or three-fourths or more the length of compatible tubes at the time of floral senescence. About 10 days after anthesis, growth of both types of pollen tubes decreases. The detached style method of pollen-tube cultivation is validated in the cultivar ‘Ace’ by seed set obtained following self-pollination in the 6- to 9-day interval and failure of seed set after either self- or cross-pollination after 9 days following anthesis. Also, in agreement with detached style data, self-pollination fails to produce seeds when attempted in bud stages or the five days following anthesis. Cross-pollinations are successful in this period. This material and technique appear well suited for study of the nature of the self-incompatibility reaction.     

Growth ofTrifolium pratense L. pollen tubes in compatible and incompatible styles of excised pistils

Summary The stigma and style portions of red clover pistils were culturedin vitro after cross- (genetically compatible) or self- (incompatible) pollination. Pollen tubes grew through styles in which they were compatible and, in some treatments, through styles which were incompatible.Boric acid, calcium nitrate and a carbohydrate in the medium, and moderate-to-cool temperatures generally enhanced pollen tube growth of compatible and incompatible matings. Several plant hormones at high concentrations inhibited pollen growth with compatible matings and did not retard the incompatibility mechanism. Application of high temperatures to the flower heads during the period of anthesis retarded the incompatibility mechanism. This temperature treatment affected the styles and not the pollen. In one of three experiments the application of relatively large amounts of pollen to the stigma rendered the incompatibility mechanism less effective.

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Zusammenfassung Narben und Griffel von Rotklee-Stempeln wurden nach Fremd- (genetisch kompatibel) oder Selbstbestäubung (inkompatibel) in vitro kultiviert. Die Pollenschläuche wuchsen durch die Griffel, mit denen sie kompatibel waren, und nach bestimmter Behandlung auch durch Griffel, mit denen sie inkompatibel waren. Ein Zusatz von Borsäure, Kalziumnitrat und einem Kohlenhydrat zum Nährmedium sowie mäßige bis kühle Temperaturen förderten im allgemeinen das Pollenschlauchwachstum der kompatiblen und inkompatiblen Kombinationen. Verschiedene Wuchsstoffe, die dem Kulturmedium zugesetzt wurden, hemmten bei höherer Konzentration das Pollenschlauchwachstum kompatibler Kombinationen, verzögerten auf der anderen Seite aber nicht den Inkompatibilitätsmechanismus. Letzterer Effekt trat ein, wenn die Blütenköpfe während der Anthesis hohen Temperaturen ausgesetzt wurden. Diese Temperaturbehandlung wirkte auf die Griffel, nicht auf den Pollen. In einem der drei Experimente führte eine Bestäubung der Narbe mit großen Pollenmengen dazu, den Inkompatibilitätsmechanismus weniger wirksam werden zu lassen.

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Cooperative research by Crops Research Division, Agricultural Research Service, U. S. Department of Agriculture and the Kentucky Agricultural Experiment Station. This paper (No. 68-3-23) is published with permission of the Director, Kentucky Agricultural Experiment Station.     

Modification of the Pollen-Stigma Interaction in Brassica oleracea by Water

The presence of a film of distilled water on the stigma surfaceof freshly opened flowers results in complete inhibition ofpollen following both incompatible and compatible pollinationsin self-incompatible (SI) genotypes of Brassica oleracea, SIgenotypes of B. campestris and one self-compatible (SC) genotypeof B. campestris. The application of water to the stigmas afterpollination also resulted in a marked reduction in pollen germinationand tube penetration. An increase in the time intervals betweenthe application of pollen onto the stigma and the water treatmentprogressively reduced this inhibition. Pollen germination wasalso completely inhibited when stigmas from freshly-opened flowersof SI B. campestris and B. oleracea genotypes were washed inwater, dried and pollinated with pollen grains of either compatibility.The ability of stigmas to induce pollen germination and tubegrowth was restored over a period, the length of which was dependenton the incompatibility (S) genotype. Stigmas of B. napus (SC)and SC mutants of SI B. campestris were found to be affectedby washing, but stigmas of a SC variety of B. campestris andthe immature stigmas from buds of B. oleracea were found tobe considerably less affected. Microscopic examination of pollenplaced on washed stigmas reveals that grains, irrespective oftheir compatibility, fail to hydrate normally. When inducedto hydrate by raising atmospheric humidity, pollen grains onwashed stigmas did germinate, but most of the tubes failed topenetrate the papillar wall and very few entered the style.It is proposed that the water treatment mobilises componentsof the pellicle which reorganize to block the activity of molecules,present in both SC and SI individuals, responsible for establishingfull contact between the pellicle and pollen grain coating. Brassica, pellicle, pollen, recognition, self-incompatibility     

Experimental insights on the function of ancillary pollen and stigma polymorphisms in plants with heteromorphic incompatibility

Most heterostylous plants possess a reciprocal arrangement of stigmas and anthers (reciprocal herkogamy), heteromorphic self‐incompatibility, and ancillary polymorphisms of pollen and stigmas. The topographical complementarity hypothesis proposes that ancillary polymorphisms function in the rejection of incompatible pollen thus promoting disassortative pollination. Here, we test this hypothesis by investigating patterns of pollen transfer and capture in populations of dimorphic Armeria maritima and A. pubigera and distylous Limonium vulgare (Plumbaginaceae), and by studying pollen adherence and germination patterns in A. maritima following controlled hand‐pollinations. Armeria lacks reciprocal herkogamy allowing the evaluation of the extent to which ancillary polymorphisms affect the composition of pollen loads. We compared the amounts of compatible and incompatible pollen on stigmas in natural populations and calculated the proficiencies of pollen transfer for each mating type. We detected disassortative pollination in each species, and mating types did not differ in compatible pollen capture, although cob stigmas captured more incompatible pollen. Controlled hand‐pollinations revealed the failure of incompatible pollen to adhere and germinate on stigmas. Our results provided evidence that, while structural in nature, pollen‐stigma dimorphisms are tightly associated with heteromorphic incompatibility and likely function to promote disassortative pollination, especially in the absence of reciprocal herkogamy.     

Changes of electric potential in pistils of Petunia hybrida Hort. and Brassica napus L. during pollination

The pattern of electric signals accompanying compatible and incompatible pollination were studied in pistils of petunia (Petunia hybrida L.) and rape (Brassica napus L). Electric potential was recorded for 4–7 hours with non-polarizable Ag/AgCl electrodes implanted into the ovary and beneath the sigma. At the end of measurements, pistils were fixed and the growth of pollen tubes was analyzed under a fluorescent microscope. Action potentials appeared in both species. In rape the potential dropped by 10 mV for few minutes after pollination regardless of the compatibility of the cross. In this species, during compatible pollination action potentials with amplitudes of 15–20 mV were recorded up to one hour after pollination. They were followed by a long lasting decrease of the potential by 10 to 50 mV. Contrary, after the self-incompatible pollination, action potentials were rare and of lower amplitudes and the potential gradually raised in comparison to the initial level. During the first hour after the compatible pollination of Petunia hybrida series of action potentials with amplitudes reaching 10–20 mV were recorded. At the time corresponding to the pollen tubes entrance to the transmitting tissue of the style, action potentials reaching up to 40 mV were followed by a steady decrease of the potential. The electric signals traveled along the style with velocity of 25 mm/s. Incompatible pollination in petunia resulted only in minor oscillation and gradual increase of the potential up to 100 mV in comparison to the initial level. The present investigation demonstrated that each phase of pollen-stigma recognition events, germination and growth of pollen tubes within the style have its characteristic pattern of electric changes which was species specific and depended on compatibility of the cross.     

INTERCELLULAR ADHESIONS IN THE POLLEN-STIGMA SYSTEM: POLLEN CAPTURE,GRAIN BINDING,AND TUBE ATTACHMENTS

Four independent pollen-stigma binding forces are differentiated after pollen contacts stigmas of Brassica oleracea var. capitata. Identification is based on their different rates and sequence of appearance during gametophyte development; on their differential occurrence after compatible and incompatible pollinations; and on their different stabilities to NaOH and hexane. The first binding force develops most rapidly, begins seconds after pollen-stigma contact, is complete within minutes, occurs on compatible or incompatible papillae, and dissociates in methanol. A second slower binding reaction begins about 15 min after contact, continues for at least 90 min (when pollen tubes emerge), results in a binding structure termed the ocreatine, develops only on compatible papillae, and is dissociated by NaOH. A third attractive mechanism binds the tip of the emerging tube to a cuticle, is detected only on incompatible papillae, and is not dissociated by NaOH or methanol. Ocreatine formation and tube development beyond the emergence stage are prevented by the incompatibility response. A fourth attraction mechanism occurs between the surfaces of papilla and elongating tubes. Reviews of physical and biochemical evidence indicate that van der Waals forces and enzymatically mediated lipid polymerization are alternatives to agglutination as mechanisms for binding male gametophyte to papilla.     

Effect of temperature on vegetative growth among isolates of Metarhizium anisopliae and M. flavoviride

Effects of temperature on vegetative growth on a semi-synthetic medium of 22 isolates of Metarhizium anisopliae and 14 isolates of M. flavoviride were determined. The majority of isolates of both species grew between 11 and 32°C; several isolates grew at 8 and 37 °C. None of the isolates grew at 40 °C. Relative growth rate, calculated from the maximum growth rate for each isolate, was significantly affected by temperature and isolate, with significant isolate * temperature interactions. The maximum absolute growth rates among the isolates ranged from 2.5 mm to 5.9 mm/day. Optimal temperatures were generally between 25 and 32 °C with several isolates exhibiting optimal growth at temperatures as high as 32 °C. Overall, relative growth rates were greater in isolates of M. anisopliae than M. flavoviride at temperatures of 25 °C or lower; conversely mean relative growth rates were greater in M. flavoviride than M. anisopliae at temperatures higher than 25 °C. However, the two most cold tolerant isolates at 8 °C were M. flavoviride and the three most heat tolerant at 35 °C were M. anisopliae. Since temperature growth responses varied considerably between isolates, strain selection according to thermal tolerance may be warranted when choosing a strain for development as a microbial control agent.This revised version was published online in October 2005 with corrections to the Cover Date.