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Developmental Patterns in Stem Primary Xylem Of August, 1967] BENZING-PRIMARY XYLEM OF RANALES. II. 813 ESAU, K. 1965a. Vascular differentiation in plants. gymnosperms. Ph.D. thesis. University of Michigan, Holt, Rinehart and Winston, New York. Ann Arbor. 1965b. Plant anatomy. 2nd ed. John Wiley MONEY, L. L., I. W. BAILEY, AND B. G. SWAMY. 1950. and Sons, Inc., New York. The morphology and relationships of the Moni­ EZELARAB, G. E., AND K. J. DORMER. 1963. The miaceae. J. Arnold Arbor. 31: 372-404. organization of the primary vascular system in the PHILIPSON, W. R., AND E. E. BALFOUR. 1963. Vascular Ranunoulaoeae. Ann. Bot. 27: 23-38. patterns in dicotyledons. III. Anatomy of the stem FAHN, A., AND I. W. BAILEY. 1957. The nodal anatomy of Calycanthaceae. Trans. Roy. Soc. Edinb. 52: and the primary vascular cylinder of the Calycan­ 517-530. thaceae. J. Arnold Arbor. 38: 107-117. QUINLAN, C. E. 1920. Contributions towards a knowl­ JACOBS, W. P., AND I. B. MORROW. 19,'')7. A quantita­ edge of the anatomy of the lower dicotyledons. III. tive study of xylem development in the vegetative Anatomy of the stem of Calycanthaceae. Trans. shoot apex of Coleus. Amer, J. Bot. 44: 823-842. Roy. Soc. Edinb. 52: 517-530. JEFFREY, E. C. 1917. The anatomy of woody plants. WETMORE, R. H., AND C. W. WARDLAW. 1951. Experi­ University of Chicago Press. mental morphogenesis in vascular plants. Ann. Rev. KUMARI, G. K. 1963. The primary vascular system of Plant Physiol. 2: 269-292. Arner. J. But. 54(7): 813-820. 1967. DEVELOP1VlENTAL PATTERNS IN STEM: PRIMARY XYLEM OF 'WOODY RANALES. II. SPECIES WITH TRILACUNAR AND MULTILACUNAR NODES1 DAVID H. BENZING Department of Botany, University of Michigan, Ann Arbor" ABSTRACT Patterns of early protoxylem strands in vegetative stems of 16 species of woody Ranales rep­ resenting nine families with trilacunar or multilacunar nodes are illustrated and discussed. The order of differentiation of first protoxylem in all examined species is basipetal. Open, protoxylem strand patterns are more common than closed patterns in these taxa and 11 other woody ran­ alean species with unilacunar nodes. Protoxylem strand patterns in woody Ranales are diverse, even among species with identical phyllotaxis and nodal anatomy. Pattern similarities of possible taxonomic significance exist, however, between some species of the Annonaceae and Myristica­ ceae, Hernandiaceae and Lauraceae, and the Canellaceae and Winteraceae. The general diversity of strand patterns in woody Ranales may reflect the high degree of evolutionary isolation among ranalean families. Results of these investigations on stem anatomy of woody Ranales indicate that the two-trace, unilacunar node is probably not primitive in the Angiospcrmao. Evidence from this study and others suggests that the primitive node in the Angiospermae, and possibly in all seed plants, is one-trace, unilacunar or trilacunar, Two-trace, unilacunar nodal anatomy of the Clerodendron type may have evolved in conjunction with the evolution of decussate phyl­ lotaxis. The protoxylem strand patterns of Illicium and/or Drirnys are probably the most primi­ tive of those described in the woody Ranales. IN A PRECEDING STUDY (Benzing, 1967) pat­ patterns are diverse and often differ among terns of protoxylem strands in vegetative shoots species possessing identical phyllotaxis. In this of 11 woody ranalean species characterized by paper patterns of protoxylem strands of 16 unilacunar nodal anatomy were described. These woody ranalean species showing trilacunar and multilacunar nodes are considered. Materials, I Received for publication 1 June 1966. This work was based on a dissertation submitted to the methods, and terminology utilized here have been Graduate School of The University of Michigan in partial discussed in the previous paper. fulfillment of the requirements for the degree Doctor of Philosophy. OBSERVATIONs-Annona muricaia, Asimina Sincere appreciation is extended to Dr. Charles B. Beck and Dr. Warren H. Wagner, Jr., for their advice during triloba, Cananga odorata (Annonaceae), My­ the course of this research and in the preparation of this ristica fragrans (Myristicaceae)-The phyllotaxis manuscript. Appreciation is also extended to the collectors and protoxylem strand pattern of vegetative mentioned in Table 1. 2 Current address: Department of Biology, Oberlin shoots of the three members of the Annonaceae College, Oberlin, Ohio. and the single member of the Myristicaceae 814 A~mRICAN JOURNAL OF BOTANY [Vol. 54 2 1 It •: 2 3 r i 3 I ., 4 U 4 5 n 4 5 1 .. n • 6I 6 4 7 4 7 8 • H 8 i Ii r~ f 4 5 • 6 4~ ~ • 2 2 4~ 3 4 I" 3 iI 4~ ) j .; 4 4~ ~ 5 . 4 , -, 5 6 n ,- •I , , : / ~I : . 6 :7 U • : IJ '" / I y~ 1/ ./ r 1 r r l f 7 . 8 August, 1967] BENZING-PRIMARY XYLEM OF RANALES. II. 815 investigated in this survey are sharply distinct in Drimys unnieri stems. Each sympodium gives from those of the ranalean species discussed in rise to both median and lateral leaf-trace strands. the previous paper (Benzing, 1967). Leaf arrange­ Leaf-trace strand divergence is exclusively cath­ ment exhibited by these four species is distichous, odic whether the ontogenetic spiral is sinistrorse and the nodal anatomy is trilacunar (Table 1). or dextrorse. The number of helicoid syrnpodia The strand patterns representing Annona muri­ agrees with the denominator of the 2/5 phyllo­ cata and Asimina triloba (Fig. 2) and Cananga tactic fraction observed for all five of these odoroia (Fig. 1) reflect the relationship between species. Three-trace, trilacunar nodal anatomy strand pattern and phyllotaxis. is common to all five species. Additional com­ In vegetative stems of all four species, two ments on the stem morphology of the Canel­ pairs of scorpioid sympodia exist; each pair pro­ laceae and the Winteraceae have been published duces median leaf-trace strands to a particular by Wilson (1960) and Bailey and Nast (1945), rank of leaves. The strand contributed by one respectively. scorpioid sympodium of a pair in Annona muri­ Figure 3 reveals the consistently repeating cata, Asimina triloba and Cananga odoraia divides nature of this strand pattern. Every fifth leaf on into two; the contribution from the other sym­ the axis is supplied by a median leaf-trace strand podium of the pair remains single. Hence, at from the same sympodial system. Lateral leaf­ departure from the vascular cylinder, the initial trace strand vascularization of each leaf is also protoxylem vascularization to the young leaf regular. Every leaf is supplied by one lateral from the central gap consists of three closely strand from the same sympodium producing the placed strands. In contrast, Myristica fragrans median strand to that leaf. The other lateral exhibits a single median leaf-trace strand which strand arises from the sympodium on the anodic results from the fusion of two unbranched strands side of the sympodium furnishing the median from a sympodial pair. This fusion product then trace strand to that leaf (see node 12, Fig. 3). passes out through the central gap as the median The length of median and lateral leaf-trace trace strand. Cananga otloraia (Fig. 1), unlike strands is irregular. Occasional lateral strands Annona muricata, Asimina triloba and Myristica fuse with other laterals and the common strand [raqrans, exhibits a consistent delay in the then fuses with a sympodium at a lower level. development of one of each pair of sympodia Of the five species of the Winteraceae and furnishing median leaf-trace strands. CaneIIaceae investigated, only Bubbia sp. ex­ The lateral leaf-trace strands in all four species hibits anastomoses between adjacent sympodia. form two groups positioned between the two pairs The resulting increase in strand number per stem of scorpioid sympodia. One lateral leaf-trace cross-section may be related to the unusually strand is furnished to every leaf by each of the large stem diameter characteristic of young twigs two lateral strand groups. In a few instances of this plant. lateral strands arise from the scorpioid sympodia, More originate from pre-existing lateral strands, IIimantandra belgraveana (Himantandraceae)­ aud the pattern is basically open. At a given Two types of leaf arrangement with correspond­ level, median leaf-trace strands show earlier ingly distinct patterns of protoxylem strands basipetal differentiation of first protoxylem ele­ were found in vegetative shoots of II. belgraveana. ments than the lateral leaf-trace strands. The strand pattern of shoots possessing distichous phyllotaxis is illustrated in Fig. 4. The strand Bubbia sp., Drimys colorata, Drimys uiinieri, pattern of other shoots from the same collection (Winteraceae), Comella alba, Warburgia uqandensis exhibiting 2/5 spiral leaf arrangement is indi­ (Canellaceae)-The .configuration of the P1'Oto­ cated in Fig. 5. Distichous shoots of II. belgraveana xylem strand pattern of Drimys uiinieri (Fig. 3) possess a grouping of median and lateral leaf­ is also characteristic of Drimys colorata, Canella trace strands. Shoots with spiral phyllotaxis alba, Warburgia uqandensis and, to a lesser extent, show an interspersion of median and lateral Bubbia sp. Therefore, a discussion of the strand leaf-trace strands. Both patterns are open and pattern of Drinujs unnieri and additional com­ lack sympodia, although occasional fusions occur ments concerning Bubbia sp, will suffice for all between lateral leaf-trace strands. The three­ five species. trace, trilacunar nodal anatomy of this species Five distinct helicoid sympodia are apparent has been described by Bailey and Smith (1942). Fig. 1-8. Solid lines depict protoxylem strands made up of mature xylem elements. Dotted lines represent strands devoid of fully differentiated xylem elements. Lines terminated by large black circles are median leaf trace xylem strands. Lines terminated by smaller black circles are lateral leaf trace xylem strands. The arrows indicate the direction of the ontogenetic spiral.-Fig. 1. Cananga odorata.-Fig. 2. Asimina triloba and Annona muricata.-Fig. 3.
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