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Salix, willow Klaus Ammann, 28. January 2007, For feedback, email [email protected] 1. Taxonomy ......................................................................................................................................................... 2 1.1. General remarks...........................................................................................................................................................2 1.2. Family Salicaceae........................................................................................................................................................2 1.2. The genus Salix ...........................................................................................................................................................8 5. Reproduction biology ..................................................................................................................................... 17 6. Risk assessment of transgenic salices............................................................................................................. 20 6.3. Risk assessment: some characteristics for transgenic trees and poplar in particular..................................................20 7. Cited literature ............................................................................................................................................... 26 Fig. 1 (Below) List of the European species of Salix according to the system of (Skvortsov, 1968), taken from (Rechinger, 1992) .................................................................................................................................................. 3 Fig. 2 The single most-parsimonious combined tree found with successive weighting. The tree has 10.271 steps (Fitch length: i.e. equal weights) with CI = 0.16 and RI = 0.38. Numbers avobe the branches are the numbers of estimated changes (ACCTRAN optimization). Underlined numbers below branches are bootstrap values; branches without an underlined number had bootstrap percentages of less than 50%. – A (left). First-branching portion of the tree, arranged with Ceratophyllaceae as the outgroup. Magnoliids form a grade composed of two major subclades (magnolid I and II) with the former sister of the eudicods. Within eudicots, ranunculids and hamamelids form a grade. The caryophyllids are sister to the asterids/rosids (for rosids, see Fig 4B). - B (right). Rosid clade. Note that the glucosinolate and nitrogen-fixing families form clades. Taxa for rbcL sequences were unabailable. + Nitrogen-fixing familiy outside the main nitrogen-fixing clade Fabaceae). From (Nandi et al., 1998) p. 153..................................................................................................................................................................... 6 Fig. 3 Dendrogram of Populus and Salix accessions, constructed from AFLP fragment similarities (Dice coefficient), with the UPGMA clustering method, and based on AFLP markers resolved by five primer combinations (EcoRI+ATA/MseI+ACAA, EcoRI+ ATA/MseI+ACAC, EcoRI+ATA/MseI+ACAG and EcoRI+ ATA/MseI+ACAT, EcoRI+AAA/MseI+ACAT). Accessions marked with an asterisk are potentially mislabeled species or hybrids (see text and Table 2). Species are marked by brackets and arrows, whereas lines group sections. Fig. 1 from (Cervera et al., 2005)............................................................................................................................................. 8 Fig. 4 Relationships of Salicaceae and putative related families based on rbcL data. 1. The single minimum length Fitch parsimony tree. Branch length (number of nucleotide substitutions) is indicated above the branches and bootstrap values (100 replicates) are indicated below them. 2. The tree obtained from neighbor-joining method. Numbers near branches are bootstrap values (100 replicates). Scale of branches indicates numbers of nucleotide substitutions per sites calculated by Kimura’s two-parameter method (Kimura, 1980). From (Azuma et al., 2000)................................................................................................................................................................ 8 Fig. 5 Relationships of Salicaceae based on rbcL data. 3. The single minimum length Fitch parsimony tree. Branch length (number of nucleotide substitutions) is indicated above the branches, and bootstrap values (100 replicates) are indicated below them. 4. The tree obtained from neighbor-joining analysis. Numbers near branches are bootstrap values (100 replicates). Scale of branches indicates numbers of nucleotide substitutions per sites calculated by Kimura’s two-parameter method (Kimura, 1980), from (Azuma et al., 2000)..................... 9 Fig. 6 Relationships of grouping in previous studies and the single minimum length Fitch parsimony tree obtained for taxa of Salicaceae based on rbcL sequences. Bootstrap values are indicated below branches. From (Azuma et al., 2000).............................................................................................................................................................. 10 Fig. 7 Plot of Salix specimens by first- and second-factor scores derived from PCA of: (a) morphology characters; (b) original 20 RAPD markers; (c) 43 RAPD markers. Filled circles represent the e-type cpDNA; hybrids misidentified in the field as S. eriocephala are indicated by squares (see text). .................................................. 12 Fig. 8 Chromosome-level reorganization of the most recent genome-wide duplication event in Populus. Common colors refer to homologous genome blocks, presumed to have arisen from the salicoid-specific genome duplication 65 Ma, shared by two chromosomes. Chromosomes are indicated by their linkage group number (I to XIX). The diagram to the left uses the same color coding and further illustrates the chimeric nature of most linkage groups, from (Tuskan et al., 2006)........................................................................................................... 13 Fig. 9 (A) The 4DTV metrics for paralogous gene pairs in Populus-Populus and Populus-Arabidopsis. Three separate genome-wide duplications events are detectable, with the most recent event contained within the Salicaceae 2 and the middle event apparently shared among the Eurosids. (B) Percent identity distributions for mutual best EST hit to Populus trichocarpa CDS, from (Tuskan et al., 2006). ........................................................................ 14 Fig. 10 The single most parsimonious bifurcating unrooted tree, based on the Wagner method, representing the phylogeny of Populus, including a branch for Salix (179-181). Plain and circled numbers correspond to accession codes (Table 2) and bootstrap values (only those above 50% are shown for main branches, grouping several species), respectively. Fig 2 from (Cervera et al., 2005). ........................................................................ 15 Fig. 11 Scatter diagram of specimens plotted by stipule length and stipule width. An asterisk (*) represents the e-type cpDNA. Circle shading indicates the rDNA genotype: white, –e/e; black, s/s; half-white and half-black, –e/s (see text). Hatched boxes indicate two standard deviations about the means for pure parents, determined by the second ML iteration. Arrows indicate specimens discussed in the text. From (Hardig et al., 2000)..................... 16 Fig. 12 Caption see above, from (Alliende & Harper, 1989) ......................................................................................... 18 Fig. 13 Caption see above, from (Alliende & Harper, 1989) ......................................................................................... 19 Fig. 19 Transgenic trees in tree physiology and biotechnology from (Herschbach & Kopriva, 2002) ........................... 20 1. Taxonomy 1.1. General remarks As an introduction, first a warning word from one of the most knowledgable plant taxonomists of Europe: Why is the taxonomy of Salix so notoriously difficult ?, (Rechinger, 1992) asks and considers Salix as the ‘crux et scandalum botanicorum’: He names several reasons: All willows are dioecious, developing male and female flowers in different individuals. Flowers in many species are precocious, the flowers appearing before the leaves. There is a tendency for the current year's terminal and past year's lateral branches to differ in indumentum, leaf shape and leaf sequence. In some species, e.g. the Capreae group in addition to S. nigricans and S. hastata, parallel variation in leaf form is frequent and has misled some authors erroneously to assume hybridisation. Hybridisation has been suspected from very early times (Smith 1805) and has been proven experimentally by Wichura (1865), Nils Heribert-Nilsson (1918), etc. Periods of over- and under-estimation of hybridisation have alternated in the past. Fertility of Salix hybrids in many cases is not at all or only slightly reduced. The lack of a fertility barrier between many species makes discrimination between certain species difficult. In hybridisation one has to distinguish between primary hybrids of scattered occurrence and widespread hybrid swarms occurring where the areas of the parents overlap. In such cases it is essential to begin with the study of variation of the parental species from areas far from their overlapping areas. 1.2. Family Salicaceae Salicaceae is a family of dioecious woody trees and shrubs with a distribution primarily in
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