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  • Chimeras and Variegation - Patterns of Deceit

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    Chimeras and Variegation_Patterns of Deceit

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    Chimeras and Variegation_Patterns of Deceit

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    148 vues12 pages

    Chimeras and Variegation - Patterns of Deceit

    Transféré par

    Mick Talbot
    Chimeras and Variegation_Patterns of Deceit

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    © All Rights Reserved
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    Chimeras and Variegation: Patterns of Deceit Department of Plant and Soil § Variegation an chimerism are phenom- ena with hoth economic and scientific impor tance for horticulture an forthe study of plant development. Yet, there remains considerable confusion in the literature regarding the mean ing of these terms and the controling mecha nisms for each phenomenon. Popular maga- ines, texts, and, ess commonly, research journals contain misuses ofthe terms “varie- tation” and “chimeras” or make gross gener alizations about variegated or chimeral plants usinga limited numberof examples, Plansae called chimeras when they ae not, or are nol fecopnized to be chimeral when they are chimeral. Frequently, itis the exception tothe ‘ule thatinitsts the confusion. In his review, [present comprehensive treatment on the ‘causes of chimerism and variegation with spe- ial emphasis on defining terms and discuss- ing the genetic and anatomical reasons why variegation and chimerism exist in higher plants, The differences and similities be- tween variegation and chimerism will be con- sidered. Finally, wil introdvce some unusual tases where variegation patterns are dificult to interpret, and demonstrate what steps of tests should be performed before assumptions are made about the cause of variegation pa tem on a particular plant WHAT IS VARIEGATION? Variegation can be defined a the presence of discrete markings of different colors on an fongan or an organism, In animals, zebra and leopards are cassicexamples. In plans, varie- gationismost frequently manifestedas stripes, blotches, and streaks, oF by differences in color between leaf (or petal) margins and the leaf (or petal) midregion. Technically, nutri tional disorders suchas iron chlorosis, or pest damage such as that caused by spider mite feeding, can lead toa variegated phenotype. In this review, however, Iwill restrict my com- ‘ments to variegation that is genetically con- trolled. Variegation types can be categorized as either cell ineage type or noncellineage type (Kirk and Tilney-Bassett, 1978). With few ered Tor publicstion 24 Sop. 1996, Accepted Foepubication 30 Nov. 1996, Publications charges ‘were deiayed.in par by grant rom the Margaret and Howard B, Bigelow Gifl © the Univ. of Massachusetts at Amherst Thank Thomas Gers fo providing Petunia geamplasm, and R. Scot Poetis for discussions conceming cel ineage vr iepation. [am grateful to Richard Munson, Thomas FH. Boyle, Mare Fower, and Susan P. MeGlew for allowing ces to some thephotographed plants The cost of publishing this paper was defrayed in part by the payment of page charges. Under posta regulations this paper therefore must be hereby marked edvertsement solely to indicate tis fact ‘Assoiae Professor. HorrScience, Vou. 32(5), Avcust 1997 Michael Marcotrigiano! exceptions, cell lineage variegation occurs genetic mosaics (individuals with cells of d ferent genotypes). The cells of one color are clonally rolted, so that each colored la streak, or patch is descended by successive cll divisions from the original cell in whch the color change occurred (Fi. I top lft and Fight) Sinoe somatic plant cells do not mi sate, cell lineage variegation patterns are re- [ated directly o cel division planes, and patch (Clone) sizeand shapeis related othe duration and rate of cell division, and, toalesser extent, todfferencesin the magnitude of cellelonga- In plants with noncel lineage variegation, alleells have the same genotype but the genes ,ponsible forthe syathesis or destruction of Piginents are expressed in only some of the ces, Thiskind of variegation is very common ciences, University of Massachusetts, Amherst, MA 01003 in both animals (eg. the black and white regions on panda bears and bald eagles) and plans [e. the leaves of Caladium bicolor fnd the stripes on watermelon (Citrus lana tus) rind). With nonce lineage variegation there iso necessary relationship between the pattem o rate of cel division and the pattern ‘of colors. Instead, the “geographic location ofthe cell, rather than the cells lineage of history, ultimately determines its phenotype (Fig, 1; bottom left and right). With noncell lineage variegation all cells have the genes necessary to make or destry'the pigment but only those in the “correct” location are “in- structed” to do so. The pater seen are the result of differential gene expression between genetically identical eels. Noncel lineage variegation pattems ean betransmitted sexually fromone generation‘ Fig. Typesofvaricpationasthey late to cel postion orcell ineage-Cellineage varices ionisdisplaye| in tap lett and op right, nonce ineapevariepaton in (baton lft) and bottom right top let) Genetic mosaic Puna Dybrida eased ty transposable genetielemen| (Dosdeman ea 14) Red sectors on white corola are directly descended rom cells that have mutated. Larger sectors were inated erie in development. (top right) Varepaton caused by somatic crossing-over. Yellowleat Nicotiana tabocin (Silo) with 4 green (susu) and white (SS) twin spot. 19 this pho, the heterozygous see spear ight gray an the homer 20s tisues white e dak aaThe spo ane ret descendants from acel ht underwent sematc rosin over. (bottom ef) Miscanthus sinensis ‘Sets (poreupine gas) Cell division daring la development in grasses proceed from meristem aie sve atthe leaf base. This horizontal banding doesnot fllow cll lineage. (bottom Tight) ‘Vasiegaton expressed sll pattern, Petania hybrid Hulahoop Blue possessing prple Nowers edged in white The "white cells are white Becase oftheir postion, dept the Tat that any are dived fom the ‘pup cells ™ the next. As with mostother traits, variegation pattems vary slightly within a population as {teractions with other genes and the environ ‘ment alter gene expression. NoncellTineage variegation paterns can be inherited simply (ea, Henny, 1982, 1983; Reisch and Watson, 1984) or complexly (Boye, 1941; Boye and Rife, 1938), and may have evolved when se- lection pressures generated pattermsthateither act as camouflage against herbivores or as pattems to atract pollinators (e., pigment pattems in flowers). Nonchimeral noncelln- age variegation is the most commen cause of variegation, PATTERN VS. NONPATTERN VARIEGATION Proquently, variegation is described as pat- temed or nonpatiered. These terms merely describe phenotype and yield litle informa: tion, Pattern variegation implies predictable, regular, repeated patter between leaves (oF flowers, or stems) on the same plant $0,€.6, ifall variegated leaves orbracts are green with white margin Fig.2), they would be consid ‘ered patterned whereas the corolla of the pet nia in Fig. 1, top lft, is nonpaterned instead displaying random splashing of color with the extent of each color being variable and ‘unpredictable fr each flower onthe plant WHAT ARE THE CAUSES OF VARIEGATION? 1. Differential gene expression. The most ‘common and most misunderstood cause of variegation is differential gene expression Expression of genes only in specific parts ofan ‘organ orn specific cll layers an give rise to this phenotype. Although a multicellular or ‘gzanism can be genetically homogeneous, ll genes are not turned on at all times in all Organs, tissues, or cells. For example, atthe ‘organ level, petals can produce pigments not produced by leaves. At the tissue level, the epidermis can produce a pigment not pro- dduced by underlying mesophyll. At the cali lar evel, cells located near the margin of an organ (e.g. petal edge) ean produce ot failto produce a pigment that cells of the same geno- type produce in a different location. There fore, positional signals that regulate gene ex- pression can determine variegation patter. Most variegated plants that come “tre 19 seed fall into thisategory. Examples include striped watermelon rinds, the commercially availablepetunia lowers with bicolored corol- las, the “faces” on pansy (Viola wittrockiana) flowers, andthe purpleringsonleavesof zonal seranium (Pelargoniven xhoriorun), 2 Leaf blisters. Ina few species, vaiega- tion manifests itself as silver spots or steaks ‘oaleaves.Thsisnotcausedby achlorophyllous tissue, but rather from an unasual develop- mental patern where, on specifi regions of the lea, cells separate from the cells below them, leaving what can best be described asa blister Fig. 3) (Hara, 1957). This is an inher- ited pattem variegation (Hoch eta, 1980) and another example of noncell lineage variega- 7 Fig, Pilea cadier (aluminum pla) n this species a ptt of “blisters” appears where the uper cll layers ae detached from the Tomer call ayers The plants ta tion, In some regions, genes responsible for allowing the cells to separate are expressed. A scientifically documentedexample of this ype of variegation isin Pilea cadien (Vaughn and Wilson, 1981), while Pulmonaria officinalis, Zebrina pendula, and maay spotted begonias (e.,Begonia maculata) appearto possess same phenotype. This type of variegation is generally patterned and, therefore, positional Signals may be involved in determining where blisters appear 3, Virus, Many viruses initiate variegation by causing nonuniform chlorosis. Tobscco mosaic virus, for example, causes a green/ chlorotic variegated phenotype on the leaves, ‘ofmany species, Generally, thesymptoms are ‘unattractive and the virus is debilitating, Yer, afew viruses add attractive variegation pat tems to either foliage or petals and do not severely impair the growth of the plant. Autilon mosaic virus causes beautiful varie ‘eaion pattersin theleaves of certain Autilon Species. The leaves of Abutilon pictum “Thompsonii are marked with bright yellow land cream patches delineated by leaf veins (Fig. 4). This vius canbe seed or graft trans- mitted (Fulton, 1968) In some hybrid tlips (Tutipa), tepals are variegated because of a Virus and the flower is streaked or striped (Fulton, 1964). In certain genotypes, the ves lative portionsof the planismay be sunted by this virus, butinothers the vegetative portions appear normal. The virus is transmitted to the daughter bulbs and the variegated phenotype is easly maintained by vegetative propaga tion. Other plants in which the variegation pattem is uncharacterized may be virusin fected. Grafting tononvariegated plants ofthe same species may indicate whether or not a virus isthe causal azent,asmost viruses trans- mit readily via the graft union 4. Genetic mosaicism. Genetic mosaicsare plants or animals in which eels of different _gorotypes coexist in a single organism, The genetic difference need not be phenotypically ‘obvious, so, ea genetic mosaic could exist for differences in the levels of a biochemical metabolite. It would not be surprising if all long-lived plants (e.., trees) are genetic mo. saics because, in the absence of a flawless system to purge cells that have spontaneously mutated, at least some cells would genetically differ from others. Patches of mutant cells would be scattered randomly throughout the ‘organism. Many bud sports of horticultural importance have arisen in this manner and have been vegetatively maintained as new cultivars, However, most mutations are not phenotypically obvious and go unnoticed Plants ean become messes duet the action of transposable genetic elements (Doodeman et al., 1984) oF spontaneous oF induced muta HorrScuence, Vor. 32(5), Auaust 1997 i 4. Leaf of Abuslm picum “Thompson an cenamental in which the variegation x cased by avr. tions in the nuclear or chloroplast genome (Tilney-Bassett, 1986). (CHIMERAS A plantchimeraisaspecifictypeof genetic ‘mosaic in which the genetically dissimilar cells are present in the shoot apical meristem, ‘where they contnve to give rise tothe cells that form the body of the plant. The arrange: ‘mentof genetically dissimilarcellsin the shoot apex is crucial to the sibility of a chimeral sate and will dictate the plan's phenotype. Several arangements are possible because shoot apices in higher plans are multicellslar and, with few exceptions, multilayered. Strict anticlinal divisions in the outer shoot apical Jayerts) give rise toa stratified meristem with 8 “tuniea-corpus” organization (Schmidt, 1924). Such divisions prevent cells in one Jayer from invading another layer. Cells inthe “tunica” layers divide antilinlly while those Jin the corpus divide in any plane. Most an: siosperms have either two or three well-de- fined apical cell layers. In most dicots, there are three apical cell layers that remain inde pendent from each other (Tilney-Bassett, 1986), whilein monocots, bothtwo-andthree Jayered apices are common (Stewart and Deemen, 1979) ‘Although the source of genetic heteroge- neity resides in the apical cell layers of the ‘meristem, one can seldom determine if plant 's chimeral by microscopie observation ofthe shoot apex. The phenotypically evident traits are generally expressed inorgans derived from themeristem rather thaninthe meristemitsef, Therefore, the chimeral composition of the shoot apex is most frequently deduced by the phenotype of the organs derived from it. In fact, the verification of the concept of chimeral plants rose from attempts at explaining what caused the white margins on the leaves HorrScience, Vot. 32(5), Aucust 1997 of variegated geraniums (Baur, 1908). (Cytochimeras—chimeras in which different apical layers possess different ploidy levels (Gatinaet al, 1940)—arethe exception, that longitudinal sections through the apex gener- ally identify chimeras, eeause apical ell size isfrequenty augmented with increasing ploidy (Stewart etal, 1972) “Three kinds of chimeras are defined by the arrangement of genetically different cells in the shoot apical meristem, The first is the sectrial chimera (Fig. 5a). With ectorial ehi- meras.a wedge of genetically unique tissue is present through all pical cell layers. Sectoral chimeras are diffcalt to identify because, in ‘mosteases,thephenolypeo genetically unique ‘als is not evident in the descendants ofall throecellayers. Indltion, neatsectorsrarely Persist, as cells within a layer tend o get “out ‘of line® with cells in other layers a cell divi- son proceeds, Sectorial chimeras would most Likely arise when mutations, ether spontane- cous or induced, occur during the early stages of embryo development when the number of

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