There are more than 80 naturally occurring relatives of cultivated tobacco (Nicotiana tabacum L.) in the genus Nicotiana. In this review, we examine how and to what extent these natural germplasm resources have been utilized in hybridization and introgression experiments over the past century. To date, more than 400 interspecific Nicotiana hybrids have been reported. We focus on individual Nicotiana species involved in interspecific hybrids with cultivated tobacco produced by sexual and asexual methods, including the recently discovered grafting method. Problems related to the hybridization of N. tabacum with other species, namely, cross-incompatibility, maternal phenotypes in hybrid offspring, interspecific incongruity, lethality of juvenile hybrids, and sterility of viable hybrids, are reviewed. Among the 58 interspecific hybrids involving N. tabacum reported thus far, 25 were also reported as somatic hybrids and two were obtained only as somatic hybrids. Thirty-six sterile sexual F1 hybrids were converted to fertile or partly fertile allopolyploids. Sixteen Nicotiana species have been deployed as a source of usable traits that were introgressed into N. tabacum, offering resistance to or tolerance of pathogens or pests. The mechanisms of introgression, such as alien addition and substitution, as well as the barriers and limitations of introgression, including erratic inheritance and adverse linkages, are discussed. Thirty-one Nicotiana species used as sources of cytoplasmic male sterility in N. tabacum have produced multiple alloplasmics; most showed negative effects of alien cytoplasm but a few have been deployed successfully in hybrid cultivars of N. tabacum.

The Use of Nicotiana Species in Tobacco Improvement

Sixteen species of Nicotiana were examined for resistance to second instar larvae of the tobacco horn-worm, Manduca sexta (L.). The tests were designed so as to discriminate between antibiosis and nonpreference. High levels of antibiosis resistance were observed in several wild species such as N. gossei and species in the Repandae section. Several other species, such as N. excelsior, showed antibiosis that appeared to be due to a different mechanism than the alkaloid-trichome exudate based resistance of the above species. The data indicate that these species of Nicotiana may be used as a new source of resistance to larvae of M. sexta.



RESUME

D'especes sauvages de Nicotiania comme nouvelle source de resistance du tabac a Manduca sexta



La resistance aux chenilles de second stade de Manduca sexta L. a ete examinee chez seize especes de Nicotiana. Les essais ont ete concus pour distinguer absence d'appetance et antibiose. Plusieurs especes sauvages comme N. gossei et les especes de la section Repandae ont presente un niveau eleve d'antibiose. D'autres especes, comme N. excelsior ont presente une antibiose vraisemblablement due a un mecanisme different de la resistance des especes precedentes provoquee par un exsudat alcaloide des trichomes. Les resultats indiquent que ces especes de Nicotiana peuvent etre utilisees comme nouvelle source de resistance aux chenilles de M. sexta.

Wild species of Nicotiana as a new source of tobacco resistance to the tobacco hornworm Manduca sexta

Four wild species of Nicotiana were examined in field experiments for antibiosis toward larvae of the tobacco hornworm, Manduca sexta (L.). The species examined were N. excelsior Black, N. gossei Domin., N. plumbaginifolia Viviani, and N. stocktonii Brandegee. Feeding, growth, and mortality rates were used as physiological indicators of plant defense compounds in a four day bioassay procedure. The magnitude of the resistance in Nicotiana stocktonii was consistently greater by all criteria measured. The mechanisms of resistance appeared to be different among the species tested. In addition, inferences on the mode of action of the resistance factors in the species and the best candidate for a breeding program are discussed.



Resume

Resistance d'especes sauvages de Nicotania aux chenilles de Manduca sexta



La resistance aux chenilles de M. sexta a ete examinee chez quatre especes sauvages: Nicotani excelsior Black, N. gossei Domin, N. plumbaginifolia Viviani, et N. stocktonii Brandagee. L'alimentation, la croissance et la mortalite pendant quatre jours d'experience ont servi d'indices physiologiques de l'action des substances chimiques de la plante. Pour tous les criteres utilises, la resistance de N. stockonii etait significativement superieure. Les mechanismes de la resistance se sont reveles differents pour les quatre especes examinees. La discussion concerne l'influence du mode d'action des facteurs de resistance et l'espece presentant le plus de potentialites pour la selection.

Antibiosis to the tobacco hornworm (Manduca sexta) in wild species of Nicotiana

Designing Tobacco Genomes for Resistance to Biotic Stresses

Experimental Introgression from Nicotiana Species to Cultivated Tobacco

In tobacco (Nicotiana tabacum L.), there is a very little scope for intervarietal gene transfer, hence interspecific hybridization utilising wild species of the genus Nicotiana possessing resistance to one or more pests, is the best solution. Eventhough N. benthamiana is resistant to tobacco caterpillar (Spodoptera litura F.), in view of its interspecific incompatibility with N. tabacum, a trispecific amphiploid (N. benthamiana×N. glutinosa×N. tabacum) was synthesised with N. glutinosa as a bridge species. Progressive elimination of N. benthamiana and N. glutinosa chromosomes was achieved through four successive back-crosses of the amphiploid with flue-cured N. tabacum varieties, resulting in plants having most of the tabacum characters and caterpillar resistance. All the plants were male-sterile and most of them possessed 48-50 mitotic chromosomes, 48 from N. tabacum and 1-2 from N. bentamiana thus indicating that the resistance factors are confined to one or two chromosomes. Based on the mortality of inoculated larvae and the extent of damage, the plants were grouped into highly resistant, moderately resistant and susceptible types.The heavy mortality of the caterpillar within 24 hours indicate that certain toxic exudates from the leaf surface may be the primary cause of death. Starvation due to “non-preference” may be another factor responsible for death of most of the remaining larvae within the next 24-240 hours. The retarded growth of a few surviving larvae, their resistant feeding and the slow pace at which they proceed from one instar to another indicate that some “antibiosis” properties may also be associated with the factors for resistance.

Interspecific Transfer of Resistance to Tobacco Caterpillar (Spodoptera litura F.) from N. benthamiana to N. tabacum.

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Interspecific Transfer of Resistance to Tobacco Caterpillar (Spodoptera litura F.) from N. benthamiana to N. tabacum.