THE GENETIC BASIS OF DIFFERENCES IN GROWTH AND BEHAVIOR OF SPECIALIST AND GENERALIST HERBIVORE SPECIES: SELECTION ON HYBRIDS OF HELIOTHIS VIRESCENS AND HELIOTHIS SUBFLEXA (LEPIDOPTERA)

Two species of moths and their hybrids were studied to determine the genetic basis of host range. One species, Heliothis virescens (HV), has a broad host range and is an agricultural pest on cotton (Malvaceae), soybean (Fabaceae), and tobacco (Solanaceae). The other species, Heliothis subflexa (HS), has a narrow host range, feeding on plants in the genus Physalis (Solanaceae). Experiments were done to determine whether the abilities to feed on cotton, soybean, and tobacco were each under separate genetic control in HV (genetically uncorrelated) or whether feeding on all three hosts was genetically correlated. By repeatedly backcrossing hybrids to HS while selecting for high larval survival and weight on soybean, loci conferring the ability to feed on soybean were moved from HV into the genetic background of the specialist, HS. After six generations of selection on soybean we tested the selected line on soybean, cotton, Physalis, and tobacco to determine if ability to feed on soybean was correlated with feeding ability on these other hosts. We found no evidence to support the hypothesis that feeding on all three hosts was correlated and controlled by a single locus. Compared to HS, the survival and weight of the selected line were significantly higher on soybean, similar on tobacco and Physalis, and slightly, but not significantly, higher on cotton. We also conducted a behavioral choice test to determine if larval preference for soybean was correlated with preference for other hosts. The selected line larvae had higher preference for soybean and cotton than HS. Preference for tobacco was equal in the selected line and in HS. These results support the trends seen in survival and growth tests where performances on soybean and cotton appeared partially correlated and performances on soybean and tobacco were uncorrelated. In conclusion, the generalist, HV, did not have a single set of loci that governed feeding on soybean, cotton and tobacco. It is likely that some of the loci governing performance and preference for soybean also contributed to performance and preference for cotton and that the soybean loci examined did not interact negatively to lower performance and preference for Physalis, the specialist's host. This approach can be applied more widely to address evolutionary questions about host range and other ecological traits.     

THE GENETIC ARCHITECTURE OF A COMPLEX ECOLOGICAL TRAIT: HOST PLANT USE IN THE SPECIALIST MOTH,HELIOTHIS SUBFLEXA

We used genetic mapping to examine the genetic architecture of differences in host plant use between two species of noctuid moths, Heliothis subflexa, a specialist on Physalis spp., and its close relative, the broad generalist H. virescens. We introgressed H. subflexa chromosomes into the H. virescens background and analyzed 1462 backcross insects. The effects of H. subflexa‐origin chromosomes were small when measured as the percent variation explained in backcross populations (0.2–5%), but were larger when considered in relation to the interspecific difference explained (1.5–165%). Most significant chromosomes had effects on more than one trait, and their effects varied between years, sexes, and genetic backgrounds. Different chromosomes could produce similar phenotypes, suggesting that the same trait might be controlled by different chromosomes in different backcross populations. It appears that many loci of small effect contribute to the use of Physalis by H. subflexa. We hypothesize that behavioral changes may have paved the way for physiological adaptation to Physalis by the generalist ancestor of H. subflexa and H. virescens.     

HYBRIDIZATION STUDIES ON THE HOST RACES OF EUROSTA SOLIDAGINIS: IMPLICATIONS FOR SYMPATRIC SPECIATION

We studied the inheritance of survival ability in host-associated populations of the tephritid fly, Eurosta solidaginis, to test predictions of sympatric speciation models. Eurosta solidaginis induces galls on two species of goldenrod, Solidago altissima and S. gigantea. The host-associated populations have been hypothesized to be host races that originated in sympatry (Craig et al. 1993). We found evidence for disruptive selection for host use, which is a critical assumption of sympatric speciation models. Each host race had higher survival rates on their host plant than on the alternative host. F₁ and backcross hybrids also had lower survival rates than the pure host-race flies on their host plant. Since assortative mating occurs due to host-plant preference (Craig et al. 1993) this would select for divergence in host preference. Low hybrid survival could have been due to strong genetic incompatibilities of the populations or due to host adaptation by each population. Strong genetic incompatibilities would result in poor survival on all host plants, while host adaptation could result in low overall survival with high hybrid survival on some host plants with particularly “benign” environments. High survival of F₁, F₂, and backcross hybrids on some plant genotypes in some years supported the host adaptation hypothesis. F₁ flies mated and oviposited normally and produced viable F₂ and backcross hybrids indicating gene flow is possible between the host races. A few flies developed and emerged on the alternative host plant. This demonstrates that genes necessary to utilize the alternative host exist in both host races. This could have facilitated the origin of one of the populations via a host shift from the ancestral host. The inheritance of survival ability appears to be an autosomal trait. We did not find evidence that survival ability was maternally influenced or sex linked.     

寄主植物对桃蛀螟生长发育及产卵选择行为的影响

为明确寄主植物对桃蛀螟生长发育及产卵选择行为的影响,利用实验种群生命表和二项产卵选择试验,研究了玉米、大豆、棉花和桃等4种寄主植物对桃蛀螟种群生长发育及产卵选择性的影响。结果表明:取食棉花的桃蛀螟幼虫存活率最低、幼虫历期最长,取食玉米的幼虫存活率最高、幼虫历期最短,取食桃和大豆的幼虫存活率和历期居于棉花处理组和玉米处理组之间;玉米处理组的桃蛀螟化蛹率、蛹重和蛹历期均为最高,棉花处理组为最低,大豆和桃处理组的这些参数均显著小于玉米处理组而大于棉花处理组;发育至成虫后,取食玉米的桃蛀螟羽化率显著高于其他3个处理组;取食桃的桃蛀螟成虫寿命(雌虫和雄虫)及个体发育历期均显著高于其他3种处理组;同时取食桃的桃蛀螟单雌产卵量最高,其次是玉米处理组,两者均显著高于大豆和棉花处理组。二项产卵选择试验结果显示,桃蛀螟雌蛾在棉花和玉米处理组、玉米和大豆处理组或棉花和大豆处理组间的落卵量差异不显著;但在包含桃的处理组中,桃蛀螟在棉花、玉米或大豆处理区的落卵量均显著高于桃处理区。上述结果表明,供试4种寄主植物中,桃蛀螟偏好在棉花、玉米和大豆上产卵,其中玉米对桃蛀螟的适合度相对较高,棉花对桃蛀螟的适合度相对较低。     

Detection of ecological hybrid inviability in a pair of sympatric phytophagous ladybird beetles (Henosepilachna spp.)

Ecological speciation is a process by which reproductive isolation evolves as the result of divergent natural selection between populations inhabiting distinct environments or exploiting alternative resources. Ecological hybrid inviability provides direct evidence for ecological speciation. To detect ecological hybrid inviability, we examined survival rates to the second instar of F1 hybrids and backcross hybrids in a pair of sympatric phytophagous ladybird beetles, Henosepilachna niponica Lewis and Henosepilachna yasutomii Katakura (Coleoptera: Coccinellidae: Epilachninae), reared on their respective host plants, thistle [Cirsium alpicola Nakai (Asteraceae)] and blue cohosh [Caulophyllum robustum Maxim. (Berberidaceae)], and on a common food plant, Japanese nightshade [Solanum japonense Nakai (Solanaceae)]. Hybrid larvae reared on leaves of the Japanese nightshade always had high rates of survival, irrespective of the crossing type of their parents, suggesting a lack of intrinsic hybrid inviability between the two species. In contrast, survival rates on thistle and blue cohosh varied greatly. On blue cohosh, the survival rate of F1 hybrids was nearly as high as that of H. yasutomii, but on thistle, survival was significantly lower than of H. niponica. Survival rates of backcross hybrids on the two host plants were intermediate between those of the parents, showing a reversed rank order of different types of backcross hybrids on the two food plant species. These results suggest that ecological hybrid inviability exists between H. niponica and H. yasutomii, although the two species do not show intrinsic hybrid inviability. Thus, our study supports the hypothesis that H. niponica and H. yasutomii underwent ecological speciation by divergent selection.     

Organogenesis and embryogenesis inHelianthus tuberosus and in the interspecific hybridHelianthus annuus ×Helianthus tuberosus

A method of plant regeneration from cotyledons ofHelianthus tuberosus, Helianthus annuus ×Helianthus tuberosus and for the backcross of the interspecific hybrids onH. annuus was developed. Induction of somatic embryogenesis and plantlet regeneration from anther culture of the interspecific hybridsH. annuus ×H. tuberosus is reported.Cotyledons were cultured on Murashige and Skoog basal medium (MS) supplemented with indole-3-acetic acid (IAA) and 6-furfurylaminopurine (kinetin) or N⁶-benzylaminopurine (BAP). Shoot regeneration occurred on most of the media tested, but the best results were obtained on media with a high concentration of cytokinins (BAP or kinetin: 4 mg l^–1) and lower concentration of auxin (IAA: 0.5–1 mg l^–1).Embryogenic callus and adventitious buds were initiated from only two anthers of the hybridH. annuus ×H. tuberosus cultured on the MS medium containing BAP (0.2 mg l^–1) and 1-naphtalenacetic acid (NAA: 0.1 mg l^–1). Prolonged culture of these embryogenic calli and buds on the original medium with successive subculture on MS basal medium without growth regulators resulted in embryo formation and shoot differentiation. The plantlets, after rooting, were established in soil.     

Model for genetic control of Heliothis virescens

Summary A model was developed which corrects and extends an earlier one proposed for the control of the tobacco budworm, Heliothis virescens (F.), through hybrid male sterility. Population suppression is effected through the release into natural populations of the backcross progeny of a hybrid between H. virescens and a related species. Thereafter, the system perpetuates itself in nature through continual backcrossing of the fertile backcross females to native H. virescens males. When the proportion of backcross hybrid females in the total population is large enough to draw off the insemination potential of the native males, the native females fail to replace themselves. The present model demonstrated that the ratio of released backcross hybrids to natural H. virescens remains constant in a closed population. Furthermore it was shown that the release ratio necessary to achieve extinction of a closed population is related to the number of females that a male can inseminate and to the population growth rate. Release ratios required to slow natural population growth and to lessen the impact damage of releases on crop plants were also examined. Effects of selection against the backcross females on the predictions of the model were explored.     

Suitability of greenbugs, cotton aphids, andHeliothis virescens eggs for development and reproduction ofOrius insidiosus

Development, fecundity, and longevity of the predator,Orius insidiosus (Say) (Heteroptera: Anthocoridae), when reared on greenbug [Schizaphis graminum (Rondani)], cotton aphid (Aphis gossypii Glover), or eggs of tobacco budworm [Heliothis virescens (F.)] with green beans or water were examined. Developmental time was shortest when predators were reared onH. virescens eggs and beans and longest when reared on cotton aphids and water. Predators were most fecund when fedH. virescens eggs. The inclusion of beans in the nymphal diet further enhanced fecundity when fed eggs. Longevity of both females and males was significantly shorter when reared on aphids than on eggs. Beans in the nymphal diet enhanced longevity of female predators only in combination with budworm eggs. FemaleO. insidiosus were largest when reared onH. virescens eggs. Addition of green beans in aphid treatments resulted in increased size ofO. insidiosus when compared to aphids and free water.     

Paternal inheritance of mitochondria and chloroplasts in Festuca pratensis-Lolium perenne intergeneric hybrids

Organelle inheritance in intergeneric hybrids of Festuca pratensis and Lolium perenne was investigated by restriction enzyme and Southern blot analyses of chloroplast DNA (cpDNA) and mitochondrial DNA (mtDNA). All F₁ hybrids exhibited maternal inheritance of both cpDNA and mtDNA. However, examination of backcross hybrids, obtained by backcrossing the intergeneric F₁ hybrids to L. Perenne, indicated that both uniparental maternal organelle inheritance and uniparental paternal organelle inheritance can occur in different backcross hybrids.     

Analyses of Two Parasitoids with Convergent Foraging Strategies

We compared the foraging strategies of two key braconid endoparasitoids of the tobacco budworm (Heliothis virescens Fab.), Cardiochiles nigriceps Vier. and Microplitis croceipes Cresson, that differ in host and habitat range but otherwise share comparable, overlapping niches. The most important host-location cues by far for both species were materials associated with damaged plants. Both species demonstrated a significant preference for volatiles released from plants damaged by H. virescens larvae over those released from undamaged tobacco and cotton plants. In choice experiments with damaged tobacco versus cotton, M. croceipes showed a significant preference for cotton plants. In contrast, C. nigriceps preferred damaged tobacco plants. Plant compounds provoked a strong response even when released from systemically induced plants (from which damaged leaves, host, and host by-products were removed). C. nigriceps appears to have a much keener ability to locate hosts over long distances than M. croceipes. This observation may be related to the highly specialized nature of this parasitoid. The possible adaptive significance of the foraging behaviors of these two parasitoids is discussed.     

Exclusion cage studies on the impact of predation on Helicoverpa armigera in cotton

The impact of predation on Helicoverpa armigera was studied in four field cage exclusion trials on cotton in Kenya. H. armigera egg cohorts were introduced inside predator‐free and open control cages, and the impact of local predator populations on the cohort was examined. Fourteen days after inoculation, exclusion cages had four times more larvae than controls, indicating a strong impact of predation. Ants and Anthocoridae were the predominant predator groups. Exclusion cages had more damaged fruiting plant parts (squares, flowers and bolls) than the control. In the absence of predators, natural mortality of H. armigera was greater as cotton matured, and is likely to be linked to the host plant condition.     

Epibiota communities of the introduced and indigenous macroalgal relatives<Emphasis Type="Italic"> Sargassum muticum</Emphasis> and<Emphasis Type="Italic"> Halidrys siliquosa</Emphasis> in Limfjorden (Denmark)

Sargassum muticum (Phaeophyceae, Fucales) has recently been introduced to Limfjorden (Denmark) where its closest relative is the indigenous Halidrys siliquosa. Previous studies have demonstrated large quantitative (canopy biomass) and qualitative (canopy persistence) differences in the habitat available to epibiota within the canopies of these two macroalgae. We therefore hypothesised that these algae would support different epibiota communities and tested this by sampling the epibiota of S. muticum and H. siliquosa on seven occasions throughout 1997 by enclosing entire thalli in mesh bags. We found 53 epibiota taxa and, with only one exception, they were all recorded on both host species. Species richness and abundance of epibiota exhibited clear seasonal variation on both host species, although epibiota biomass was seasonally constant on H. siliquosa but not on S. muticum. These patterns were consistent with the different life histories of the host species. There was a weakly negative correlation between thallus size and epibiota biomass for both host species. When taking species-specific seasonal variation in thallus size into consideration, S. muticum and H. siliquosa were found to support significantly different epibiota biomasses. Multivariate analyses showed that epibiota community structure was different, although highly overlapping, between the two species, whereas there was an almost parallel temporal development in epibiota community structure. We conclude that it is unlikely that the introduction of S. muticum to Limfjorden has caused major changes in local epibiota community structure. However, the standing stock of epibiota is likely to have increased.Communicated by H.-D. Franke     

Natural hybridization and gene flow between twotridentiger gobies in lake hinuma (ibaraki prefecture, japan)

Of 851 specimens ofTridentiger obscurus andt. brevispinis collected from Lake Hinuma (Ibaraki Prefecture, Japan) from July 1996 to February 1998, 49 (5.8%) comprised F₁ hybrids and backcross progeny of the two species. Since the mitochondrial DNA haplotypes of the F₁ hybrids reflected those ofT. brevispinis, most instances of hybridization are thought to have occurred between maleT. obscurus and femaleT. brevispinis. Although both allozyme and mitochondrial DNA analyses indicated backcrossing and introgression of mitochondrial DNA, the frequency of backcross progeny was relatively low, suggesting the existence of a natural selection to backrossing.     

Host specificity ofPterolonche inspersa [Lep.: Pterolonchidae] and its potential as a biological control agent forCentaurea diffusa,diffuse knapweed,andC. Maculosa,spotted knapweed

In 1981 the United States Department of Agriculture (USDA) Biological Control of Weeds Laboratory (BCWL, Rome, Italy) started investigating the host specificity ofP. inspersa as a candidate for biological control ofC. diffusa andC. maculosa, and in 1985 the investigation was completed at the USDA Biological Control of Weeds Laboratory at Albany, California. Thirty five species of plants in the familyAsteraceae and 23 other species in 8 related families were tested. Larval survival beyond 1st instar occurred only onCentaurea spp. except for 1 larva found inCnicus benedictus L.P. inspersa is stenophagous onCentaurea, therefore, a good candidate for introduction into North America.      

Hybridization between Triticum aestivum L. and Agropyron michnoi Roshev.

Summary Intergeneric hybrids between Triticum aestivum cv Chinese Spring (2n=6x=42, AABBDD) and Agropyron michnoi Roshev. (2n=4x=28, PPPP) were obtained by embryo culture. Their spike characteristics were similar to those of common wheat but, unlike their parents, they were long-awned. The average meiotic chromosome pairing at MI of F₁ hybrids was: 6.39 I +3.75 rodII+8.64 ringII+0.81 III+0.30 IV+0.04 V, the bivalent and multivalent formation of which was much higher than expected from the genomic formulae. It is especially worthwhile to note that the F₁ hybrids were self-fertile, self set being 0.15%, and seeds were easily obtained from the backcross of f₁ plants with hexaploid and tetraploid wheats; here the seed set was more than 20.0%. The polyploid taxa and the position of A. Michnoi in Agropyron are discussed.     

Cytogenetic and genomic relationships ofThinopyrum elongatum with twoPsathyrostachys species and withLeymus secalinus (Poaceae)

Intergeneric hybridizations were made betweenT. elongatum, and twoPsathyrostachys and fiveLeymus species. The seed set obtained onT. elongatum ×Leymus hybrids ranged from 5.65% to 20.00%, depending onLeymus species. The seed set obtained onT. elongatum ×Psathyrostachys hybrids ranged from 16.07% to 19.70%. Meiotic pairing at metaphase-I in JN diploid hybrids ofT. elongatum ×Psathyrostachys species revealed a very low level homology between the basic J and N genomes, and further demonstrated that the two genomes are quite diverged. Chromosome pairing in theT. elongatum ×Leymus secalinus hybrid averaged 15.19 univalents + 2.62 rod bivalents + 0.26 ring bivalents + 0.02 trivalents, suggesting that the partial J^e chromosomes ofT. elongatum has homology withLeymus secalinus genomes.L. secalinus might have 3–4 chromosomes originating from J^e genome.     

The biology of two Eretmocerus spp. (Haldeman) and three Encarsia spp. Forster and their potential as biological control agents of Bemisia tabaci biotype B in Australia

The performance, as measured by daily rate of parasitism and total parasitism, of five aphelinid species found in Australia parasitising Bemisia tabaci were compared on cotton, hibiscus, rockmelon, soybean and tomato. Two Eretmocerusspp., both indigenous to Australia, gave the highest levels of parasitism on each of the plant host species tested. The tritrophic interactions between B. tabaci, host plant species and Eret. mundus(Australian parthenogenetic form) (APF) were also examined. In general, more whiteflies were parasitised when cotton was the source host or rockmelon the test host. Parasitism was always low when tomato was either the source or test host. When parasitoids were transferred from rockmelon to cotton, parasitism declined. In contrast, parasitism increased when parasitoids were transferred from cotton to rockmelon. Parasitism also increased when parasitoids were transferred from soybean to rockmelon, yet failed to do so when shifted from soybean to cotton despite cotton normally being a better host. However, when parasitoids were transferred from cotton to soybean there was a marked increase in parasitism. Possible causes are discussed. The field cage trial demonstrated that parasitism by both Eretmocerus spp. increased with increasing whitefly density. Further, the increase in parasitism was not due to the presence of more parasitoids as neither the parasitoid-whitefly ratio nor the total number of parasitoids present had a significant effect on parasitism. The combination of the two species, gave similar levels of parasitism to that achieved by Eret. mundus(APF) alone. Subsequent identification of the emerged individuals indicated that over 50% of the parasitism was due to this species suggesting that it out-competed Eret. queenslandensis. Despite this competition, there was no evidence that overall control was compromised.     

Host-associated fitness variation in a seed beetle (Coleoptera: Bruchidae): evidence for local adaptation to a poor quality host

The geographic distributions of many generalist herbivores differ from those of their host plants, such that they experience coarse-grained spatial variation in natural selection on characters influencing adaptation to host plants. Thus, populations differing in host use are expected to differ in their ability to survive and grow on these host plants. We examine host-associated variation in larval performance (survivorship, development time, and adult body weight) and oviposition preference, within and between two populations ofStator limbatus (Coleoptera: Bruchidae) that differ in the hosts available to them in nature. In one population,Acacia greggii (Fabaceae: Mimosoideae) andCercidium microphyllum (Fabaceae: Caesalpininoideae) are each abundant, while in the second population onlyC. floridum andC. microphyllum are present. In both populations, egg-to-adult survivorship was less than 50% onC. floridum, while survivorship was greater than 90% onA. greggii. Most of the mortality onC. floridum occurred as larvae were burrowing through the seed coat; very low mortality occurred during penetration of the seed coat ofA. greggii. Significant variation was present between populations, and among families (within populations), in survivorship and egg-to-adult development time onC. floridum; beetles restricted toCercidium in nature, without access toC. floridum, survived better and developed faster onC. floridum than beetles that had access toA. greggii. Large host effects on body size were detected for female offspring: females reared onA. greggii were larger than those reared onC. floridum, whereas male offspring wee approximately the same size regardless of rearing host. Trade-offs between performance onC. floridum andC. floridum were not detected in this experiment. Instead, our data indicate that development time and survivorship onC. floridum may be largely independent of development time and survivorship onA. greggii. Patterns of oviposition preference corresponded to the observed patterns of host suitability: in laboratory preference tests, beetles with access toA. greggii in nature tended to prefer this host more than beetles without access to this host in nature.     

Inflorescence culture of wheat-Agropyron hybrids: Callus induction,plant regeneration,and potential in overcoming sterility barriers

Segments of young inflorescences of Triticum aestivum cv. Chinese Spring (CS), its F₁ hybrids with Agropyron trachycaulum and A. scirpeum and backcross derivatives with A. yezoense, A. intermedium and A. junceum, and of a A. yezoense x T. aestivum cv. Wichita hybrid were cultured. Different parts of young spikelets of A. trachycaulum x CS F₁ and A. yezoense x Wichita F₁ 's were also cultured. Percent callus induction was lower in wheat than in the wheat-Agropyron hybrids or backcross derivatives. Percent callus induction from different organs in both hybrids was in the descending order of whole spikelet, spikelet without glumes, rachis, and glumes. No plants could be regenerated from calli of wheat and backcross derivatives except those of CS x A. intermedium combination. Callus induction in hybrids varied from 54 to 84% and plant regeneration from 14 to 31%. The regenerants required no vernalization. Variants including one with top-dense spikes and another with elongated spikelets were recovered. Out of eight A. trachycaulm x CS hybrid regenerants, one had anthers and stigma as opposed to neutral flowers of the original hybrid.     

Combining cry1Ac with QTL alleles from PI 229358 to improve soybean resistance to lepidopteran pests

A QTL conditioning corn earworm resistance in soybean PI 229358 and asynthetic Bacillus thuringiensis cry1Ac transgene from therecurrent parent Jack-Bt were pyramided intoBC₂F₃ plants by marker-assisted selection. Segregatingindividuals were genotyped at SSR markers linked to an anitbiosis/antixenosisQTL on linkage group M, and were tested for the presence ofcry1Ac. Marker-assisted selection was used during andafter the two backcrosses to develop a series of BC₂F₃plants with or without the crylAc transgene and the QTLconditioning for resistance BC₂F₃ plants that werehomozygous for parental alleles at markers on LG M, and whicheither had or lacked cry1Ac, were assigned to one of fourpossible genotype classes. These plants were used in no-choice, detached leaffeeding bioassays with corn earworm and soybean looper larvae (Lepidoptera:Noctuidae) to evaluate the relative antibiosis in the different genotypeclasses. Resistance was measured as larval weight gain and degree of foliageconsumption. Few larvae of either species survived on leaves expressing theCry1Ac protein. Though not as great as the effect of Cry1Ac, the PI229358-derived LG M QTL also had a detrimental effect on larval weights of bothpest species, and on defoliation by corn earworm, but did not reduce defoliation bysoybean looper. Weights of soybean looper larvae fed foliage from transgenicplants with the PI-derived QTL were lower than those of larvae fed transgenictissue with the corresponding Jack chromosomal segment. This work demonstratesthe usefulness of SSRs for marker-assisted selection in soybean, and shows thatcombining transgene-and QTL-mediated resistance to lepidopteran pests may be aviable strategy for insect control.