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Lateral Meristems, Successive Cambia and Their Products: a Reinterpretation Based on Roots and Stems of Nyctaginaceae

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Lateral Meristems, Successive Cambia and Their Products: a Reinterpretation Based on Roots and Stems of Nyctaginaceae Botanical Journal of the Linnean Society, 2004, 146 129— 143. With 24 figure’ Lateral meristems, successive cambia and their products: a reinterpretation based on roots and stems of Nyctaginaceae SHERWIN CARLQUIST ns Santa Barbara Botanic Garden, 1212 Mission Canyon Road, Santa Barbara, CA 93105, USA Received January 2003; accepted for publication April 2004 pro Anatomical studies of stems and roots with lateral growth from eight species of seven genera of Nyctaginaceae vide material for analysis of meristematic activity and histological products of that activity. Ideas about occurrence dif of meristems that achieve lateral growth in Nyctaginaceae are reviewed. The interpretation offered for the family fers from those of other workers, although the new interpretation is clearly implicit in Solereder’s figure of secondary growth in Pisonia. A lateral meristem produces secondary cortex to the outside. To the inside, it produces conjunctive succes tissue (both parenchymatous and fibrous), true rays (except in Bougainvillea and Heimerliodendron) and a vascular cam sion of vascular canibia. As each vascular cambium is produced, the lateral meristem outside of the less bium tends to become quiescent, returning to activity when the vascular cambium internal to it has become in active. Quiescence of the lateral meristem at these points coordinates the amount of tissue produced to the inside Bougainvillea zones without vascular cambia with that produced in zones with cambia. Heimerliodendron is rayless; do not has minimal differentiation between conjunctive tissue parenchyrna and ray parenchyina. Vascular cambia produce rays in Nyctaginaceae (lateral meristems do), although vascular cambia produce vessels, axial parenchyma of and sometimes fibres to the inside and indefinite amounts of secondary phloem to the outside (earlier increments readily phloem are crushed). Conjunctive tissue is held to have both parenchymatous and fibrous components and is distinguished from products of vascular cambia. Nonbordered perforation plates characterize Nyctaginaceae, as they that of do many families of Caryophyllales. The anatomical plan of Heimerliodendron is markedly different from Pisonia s. s., and Pisonia sect. Prismatocarpus (which includes Heimerliodendron in some treatments) may merit except generic recognition. The lateral meristems of all Nyctaginaceae studied are storied (stratified), as are products for fibres, which undergo such extensive intrusive growth that a storied pattern is not achieved at maturity. The families tenns ‘included phloem’ and ‘interxylary phloem’ must be abandoned for descriptions of vascular tissue in 2004, of the order Caryophyllales. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 146, 129—143. — — — lateral growth ADDITIONAL KEYWORDS: cambial variants — Caryophyilales conjunctive tissue vascular cambia — wood anatomy. INTRODUCTION mechanisms that have produced these patterns. lateral thickness may . Nyctaginaceae that increase in anatomists who have dealt with• Nyctaginaceae Plant • be said to have successive cambia (although some (see Gregory, 1994) have offered satisfactory descnp authors reject that term in Nyctaginaceae). However, tions of the ground plans of stems and roots, and which they . the source of these cambia and the way in - An provided accurate accounts of the histology. act have been subject to diverse and controversial however, . understanding of these ground plans, interpretations. The apparent diversity of ground • . ontogenetic requires appropnate knowledge of the plans in Nyctaginaceae offers opportunity, as well as difficulty in any attempt at resolution, of interpreting the full range of meristematic activity in axes of the A relatively large grouping of genera and spe *Ad&g for correspondence: 4539 Via Huerto, Santa Barbara, family. CA 93110-2323, USA. E-mail: s.carlg&st@’verizcn.net cies has been assembled here in comparison to other 129—143 129 © 2004 The L,innean Society of London, Botanical Journal of the Linnean Society, 2004, 146, 130 S. CARLQUIST studies on meristematic activity in the family, studies The above interpretations all have several plausible that have usually focused on a single genus. The diver features, especially where the species treated by the sity of meristematic manifestations and the diversity authors are concerned. All of these studies have the of products of these meristems in Nyctaginaceae disadvantage of treating only a single genus, and none relate, in my opinion, to variations on a single plan. of them include the distinctive pattern represented by Evolution of entirely disparate phenomena for achiev Neea, Quapira, Pisonia as. and Torrubia. The techni ing lateral growth in plants with similar appearing cal difficulties in sectioning woods of these genera may axes seems unlikely. A comparative method is there have delayed understanding of meristematic activity fore followed here. Compilation of detailed wood data in lateral growth of stems of the family. A summary is not deemed justified because the sampling is not headed Wood’ was presented by Metcalfe & Chalk extensive and because data based on different organs (1950) in their account of Nyctaginaceae. Metcalfe and of widely different ages are not applicable to compar Chalk called parenchymatous background tissue ‘con ative studies elsewhere in the order Caryophyllales. junctive tissue’, but the fibrous background tissue was The meristematic phenomena shown by Nyctagi termed ‘wood’ and thus they used the term ‘interxylary naceae may well be of pervasive importance within the phloem’ for phloem strands. Metcalfe & Chalk (1950) order. Just as entirely disparate methods of cambial used the term ‘rays’ in some genera of Nyctaginaceae, activity are not to be expected within Nyctaginaceae, a but ‘radial conjunctive tissue’ in others. The terms single basic plan (with variations) related to the phe used by various authors reflect their concepts for the nomenon of successive cambia seems likely present origin of the mature tissues that they discuss. throughout Caryophyllales. In turn, the interpreta Although all of the authors cited attempted to present tions offered for patterns in Nyctaginaceae seem, at accurate descriptions of histology, none of them have, present, applicable to families as diverse as Convolvu in my opinion, presented interpretations for lateral laceae and Gnetaceae. growth mechanisms that are entirely accurate, and a Some introduction to the nature of disagreements is reinterpretation of the phenomenon is therefore necessary as a background for the present observa needed. tions. Esau & Cheadle (1969), studying Bougainvillea I am hoping that by offering a survey of axes of var spectabilis, favoured the idea of a succession of cambia ious Nyctaginaceae, using methods that show both which ‘arise in centrifugal order .. each originating meristems and mature tissues clearly, the nature of among the derivatives of the preceding cambium. meristematic action can be clarified. In addition, I am Such cambial layers function bidirectionally, produc influenced by my eariier studies on anatomy of Caryo ing xylem towards the inside of the axis and phloem phyllales with successive cambia (Carlquist, 1995, towards the outside.’ In addition, they believed that 1997, l999a, 1999b, 1999c, 2000a, 2000b, 2000c, 2002, ‘conjunctive tissue and xylem fibres’ are formed 2003). The summary of Gibson (1994) has also been ‘toward the inside’. This view contrasts sharply with helpful. Above all, note should be taken that the pat that of Studholme & Philipson (1966), who worked tern of secondary activity in the stem of Pisonia illus with Heimerliodendron brunonianum. Studholme and trated by Solereder et al. (1908) and copied by Philipson claimed that there is a ‘region of maximum Metcalfe & Chalk (1950) has never been explained in meristematic activity. which progresses regularly ontogenetic terms. The concepts in the present paper outwards’, apparently by ceasing to function internal may be considered overdue explanations for this pat to phloem but then by forming outside of the phloem: tern, as well as for others in Nyctaginaceae. ‘as the development of the phloem strands proceeds, I do not believe that any useful purpose is served by the cambial cells external to it can be seen to become having two terminologies for anatomy of dicotyledons meristematically more active, so that the phloem with cambial variants: one for those knowledgeable appears to be enclosed within two arcs of the most about these plants and one for those unfamiliar with active zone. Finally, the active zone lies entirely out them. Consequently, a terminology that is accessible side the phloem.’ Stevenson & Popham (1973), work to both kinds of workers is attempted here. The term ing with Bougainvillea spectabilis, claimed a single ‘successive cambia’, used by Schenck (1893), Pfeiffer unidirectional meristem which gives rise to ‘desmogen (1926), and others can readily be applied to dicotyle strands’ (term used earlier by Pfeiffer, 1926), a concept don axes in which one sees a series of vascular strands allied to procambial strands. The desmogen strands or bands, each with both secondary xylem and second are claimed to be embedded in a background of con ary phloem. Likewise, the term ‘conjunctive tissue’ can junctive tissue. Only after phloem and xylem differen easily be used for the ground tissue in which the sec tiate from cells in these strands does a (vascular) ondary xylem and phloem are embedded (secondary
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