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Micropropagation, Genetic Engineering, and Molecular Biology Expression Vectors Containing the Gene(S) and Promoter(S) of Populus

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Micropropagation, Genetic Engineering, and Molecular Biology Expression Vectors Containing the Gene(S) and Promoter(S) of Populus This file was created by scanning the printed publication. Errors identified by the software have been corrected; however, some errors may remain. Chapter 11 Growth and Development Alteration in Transgenic Populus: Status and Potential Applications1 Bjorn Sundberg, Hannele Tuominen, Ove Nilsson, Thomas Moritz, c. H. Anthony Little, Goran Sandberg, and Olof Olsson Introduction Using Transgenic Populus to Study Growth and Development With the development of gene-transfer techniques appli­ cable to forest tree species, genetic engineering is becoming in Woody Species an alternative to traditional tree breeding. To date, routine transformation methods for several hardwood species, par­ The best model system for understanding the genetic ticularly Populus and Eucalyptus, have been established and and physiological control of tree growth and wood for­ promising advances have occurred in the development ~f mation is a perennial species containing a vascular cam­ transformation protocols for conifers (Olarest et al. 1996; Ellis bium, which is the meristem that produces secondary et al. 1996; Jouanin et al. 1993; Walters et al. 1995). Rapid xylem and phloem. Presently, Populus is the preferred tree­ progress in transformation technology makes it possible to model system because it has several useful fea.tures. develop genetic engineering tools that modify economically Populus has a small genome, approximately 5 x lOS base tractable parameters related to growth and yield in tree spe­ pairs (bp ), which encourages molecular mapping, library cies. Such work will also increase our understanding of ge­ screening, and rescue cloning. Saturated genetic maps are netic and physiological regulation of growth and already constructed for several Populus spp. (Bradshaw et development in woody species. al. 1994; Cervera et al. 1996). Moreover, Populus can beef­ Several hybrid aspen lines with phenotypes modified ficiently transformed and regenerated, and it grows rap­ by genetic engineering were produced and characterized idly. Thus, isolated genes inserted into proper vectors can in our laboratory. Their existence shows, unequivocally, easily be introduced into Populus, readily producing that tree growth and development are alterable by genetic transgenic plants. engineering. Some modified phenotypes are described in The major disadvantage of the Populus model system is this chapter. We also briefly discuss how genetic engineer­ that controlled crosses are time consuming and difficult to ing might be used to generate trees with properties, such perform. Consequently, expression of introduced genes in as modified wood structure, which are desirable to the the F generation remains undetermined. To date, all physi­ forest industry. 2 ological and phenotypic investigations involving transgenic Populus have been restricted to the primary transformant. However, as research on Populus increases, novel schemes will be developed to induce early flower­ ing by physiological, biochemical, or genetic manipulation. As discussed below, the potential for genetic manipula­ tion has been shown by the expression of the Arabidopsis LEAFY (LFY) gene in a hybrid aspen. 1 Klopfenstein, N.B.; Chun, Y. W.; Kim, M.-S.; Ahuja, M.A., eds. We used a hybrid aspen, Populus tremula x P. tremuloides, Dillon, M.C.; Carman, R.C.; Eskew, L.G., tech. eds. 1997. as a model system for genetic engineering experiments. Micropropagation, genetic engineering, and molecular biology Expression vectors containing the gene(s) and promoter(s) of Populus. Gen. Tech. Rep. RM-GTR-297. Fort Collins, CO: of interest were introduced by electroporation into the C58 U.S. Department of Agriculture, Forest Service, Rocky Mountain Agrobacterium tumefaciens strain GV3101 (pMP90RK). To Research Station. 326 p. move the transfer DNA (T-DNA) from these vectors to the 74 Growth and Development Alteration in Transgenic Populus: Status and Potential Applications Populus genome, sterile, internodal stem segments were cocultivated with the Agrobacterium for 2 days on solid Murashige and Skoog (MS) medium (Murashige and Skoog 1962). Cocultivated explants were transferred to new media containing plant growth substances to initiate shoot formation and antibiotics to select for transformants and eliminate remaining Agrobacterium cells (Nilsson et a!. 1992). Using this protocol, transformation frequency was typically about 20 percent, regeneration success was higher than 95 percent, and rooted plants 7 to 10 em tall were obtained after about 4 months in sterile culture. Transgenic plantlets were either placed in pots and cultured in a green­ house or placed in a controlled environment chamber (fig­ ure 1). For the first 7 to 10 days of culture, plants are covered with a plastic bag for acclimation to decreased humidity. Gene expression, and therefore phenotype, is signifi­ cantly affected by environmental conditions such as light quality and quantity, nutrient availability, and tempera­ ture. To compare phenotypes from different transforma­ tion events, plants must be cultured under controlled and reproducible environmental conditions, especia lly when performing experiments at different times of the year. For critical evaluation of transgenic phenotypes, we used con­ trolled environment chambers and potted the plants in mineral wool. This potting medium was used for a stable and defined nutrient supply through daily surplus water­ ing with an optimal nutrient solution (Ingestad 1970). These culture conditions induced rapid growth; about 1 leaf primordia was produced every 2 days in wild-type plants, and a plant 1 m tall was produced after about 6 weeks in the growth chamber. The Populus model system enables the function of genes isolated from any organism to be evaluated in a tree spe­ cies by transgene expression. Alternatively, the correspond­ ing endogenous Populus gene can be cloned and used for transformation in either sense (overproduction of the gene product) or antisense (suppression of the gene expression) orientation. Figure 1. Regeneration and culture of hybrid aspen plants. A) Internodal stem segments. B) Shoots regen­ erated on stem segments. C) Shoot proliferation on cytokinin-rich medium. D) Initial culture of potted plants under plastic bags. E) Plants Transgenic Populus With Altered cultured under controlled environment conditions in a climate chamber. F) Plants cultured in a Growth and Development greenhouse. Transgenic Hybrid Aspen Expressing the Arabidopsis LEAFY Gene flowers grow (Huala and Sussex 1992; Schultz and Haughn Knowledge of gene function in a distantly related model sys­ 1991; Weigel eta!. 1992), which shows that LFY is necessary tem, such as Arabidopsis, can be directly applied to a tree species. for normal flower initiation in that species. Furthermore, LFY This was recently shown by expression of the Arabidopsis gene is the first gene expressed in the flower primordium, and LEAFY (LFY) in hybrid aspen to promote precocious fl owering LFY transcription is detected in the predicted positions of (Weigel and Nilsson 1995). Arabidopsis plants with loss­ the flower primordia before any other signs of primordium of-function mutations in LFY have shoots where typically formation are evident (Weigel eta!. 1992). 75 USDA Forest Service Gen. Tech. Rep. RM-GTR-297. 1997. Section II Transformation and Foreign Gene Expression Figure 2. Typical phenotypes of hybrid aspen transformants. A) and B) Plants expressing the Arabidopsis LFY gene. Note flowers formed by the axillary (A) (Weigel and Nilsson 1995) and apical (B) meristems. C) and E) Plants express­ ing the Agrobacterium tumefaciens T-DNA IAA-biosynthesis genes. Note the slower growth, smaller leaves and, after decapitation, inhibited axillary bud outgrowth in transformants (C-left, E-right) compared with wild-type plants (C-right, E-left). D) Plants expressing the oat phyA gene. Note the short internodes. F) and G) Plants express­ ing the Agrobacterium rhizogenes T-DNA ro/C gene. Note the bushy phenotype (F) and stem fasciation (G) (Nilsson et al. 1996b). These findings suggest that LFY is the "main switch" for the fl ower formation program, and that it might be necessary and su fficient to ind uce flowering. This func­ tion of LFY was shown by fusing the LFY coding region to the strong cauliflower mosaic virus (CaMV) 355 pro­ moter and tran sforming the 355-LFY construct into Arabidopsis (Weigel and Nilsson 1995). The transgenic 355- LFYplants flowered earlier than their wild-type counter­ parts because the meristem in the leaf axil, which norma lly produces a secondary shoot, produced a single flower. Eventuall y, the apical meristem of the primary shoot fo rmed a termina l flower. However, because Arabidopsis natu rally fl owers very early, a fter about 3 weeks under inductive long-d ay conditions, the induced difference in flowering time was not very dramatic. A more stringent test of the flower-inducing capability of LFY and for the conservation of LFY function between species was the introduction of the same 355-LFY con­ struct into a hybrid aspen whose parental species take 8 to 20 years to flower under natural conditions (Schreiner 1974}. Rem arkably, the 355-LFY hybrid aspen transformants fl owered after only a few months in tissue culture (Weigel and 1 ilsson 1995) (figure 2A, 28). Transformants with high LFYexpression produced a soli­ ta ry flower in the axil of a few leaves before the apical meristem of the shoot was consumed in the formation of an aberrant te rmina l fl ower. Transformants with low LFY expression could be rooted and subsequently were transferred to the greenhouse. Some of these p lants also formed a single fl ower with normal appearance in the leaf axils, bu t it remains unclear whether these fl owers are fe rtile. Thus, the phenotype of hybrid aspen p lants that s trong ly express 355-LFY is similar to ana logous Arabidopsis transformants. This shows that the fu nction of the LEAFY protein is highly conserved between unre­ lated d icot species and raises the possibility that the Arn~idopsis LFY can be used to induce early flowering in a w1de range of tree species. This range of tree speci'es may incl ude conifers as Douglas-fir contains a gene that shows high homology with Arnbidopsis LFY (S.
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