Plant Regeneration from Protoplasts: a Literature Review

Bot. Neerl. March 1-23 Acta 38(1), 1989, p.

Plant regeneration from protoplasts: a literature review

S. Roest and L.J.W. Gilissen

Research Institute Ital, P.O. Box 48, 6700 AA Wageningen, The Netherlands

CONTENTS

Introduction 1

Plant species 2

Protoplast regeneration 3

Plant factors 10

External factors 12

Conclusions and perspectives 13

Key-words: plant regeneration, protoplasts.

INTRODUCTION

be isolated in from different tissues and Protoplasts can large quantities plant organs.

have shown the cell walls and They ability to synthesize new to regeneratecomplete plants when culturedin the appropriate media.Therefore, protoplasts are regarded as totipotent cells, which are well suited for fundamental studies on the one hand, and for genetic manipulation on the other. For crop improvement through genetic manipulation techniques, such as somatic hybridization, micro-injection and electroporation, plant regeneration from protoplasts is of essential significance.

As Af Klercker succeeded in the mechanical isolation early as 1892, of relatively low numbers of protoplasts of Stratiotes aloïdes by sectioning plasmolysed tissue. The isolation of of protoplasts Lycopersicon esculentum, using cell wall degrading enzymes

(Cocking 1960), was a great break-through in protoplast research. The enzymatic treatment enabledthe routine isolation ofrelatively high numbers of protoplasts fromalmost every organand tissue ofa diversity ofplant species. In the period between 1960and 1970, due to the combined application of cytokinins and auxins, cultured protoplasts showed the ability to form cell walls and to undergo cell divisions leading to colony formation.In

for the first from achieved 1971, time, plant regeneration protoplasts was in tobacco

(Nagata & Takebe 1971; Takebe et al. 1971). Subsequently, regeneration has been described from protoplasts of a great diversity of plant species, especially those that belong to the Solanaceae.

Binding (1985) and Maheshwari et al. (1986) have listed about 100 plant species for which successful regeneration procedures have been described. Now this number has increased more thantwofold, which justifies the publication ofan up-to-date bibliography ofall known species. In addition, plant factors and external factors, which proved to be of great importance for successful regeneration, are briefly commented on. In addition, examples ofrecently developed protoplast culture techniques are described. Finally, some problems that hamper the use of protoplasts in genetic manipulation studies are discussed and attention is given to some techniques that can be used to provide more knowledge on fundamentalaspects of protoplast regeneration.

1 2 S. ROEST AND L. J. W. GILISSEN

PLANT SPECIES

of 212 of cell From protoplasts higher plant species, representing 96 genera 31 families,

colonies and calli have been developed, which regenerated into embryo-like structures,

embryoids or shoots (Table 1). Normally, when cultured on appropriate media, these

structures are able to develop into plants. Plant species that did not yield embryoids or

shoots from protoplast-derived colonies, or that regenerated into roots only, are not

included in the list. The species are grouped according to their systematic taxa in alpha- betical order. In general, the first publications are presented in which the regeneration

for procedure a given plant species is described. From 1970until 1987, the yearly number of new plant species, for which details on the regeneration from protoplasts have been

published, is presented in Fig. 1. Obviously, during the last few years, these numbers have

increasedand for the first of rapidly 9 months 198828 new plant species can be added(not

included in Fig. 1).

Fig. I. The number of higher plant species for which regenerationfrom protoplasts has been published yearly between 1970 and 1987.

Plant species of the Solanaceae appear to be very responsive. Until now, 67 species (±30% of all listed species) of this family have demonstrated regeneration potential.

Within like the Solanaceae, including commercially important genera Lycopersicon,

Petunia and Solanum regeneration is almost exclusively achieved by Nicotiana, , plant adventitious shoot formation.

the last substantial has been made the During 5 years, progress concerning regeneration from protoplasts of monocotyledonous and dicotyledonous species, representing several

Gramineae, Papilionaceae and woody crops, that were considered to be recalcitrant in PLANT REGENERATION FROM PROTOPLASTS 3

Table 1. (a) List of abbreviations used in Table 1(b)

Donor tissue Culture technique Regenerant development

B - C = callus bead type C =callus, cell colony

D E = Co= cotyledon, hypocotyl = droplet (20-200 pi) somatic embryo, embryo-

like structure

L = P leaf, shoot apex L =liquid medium =plant(let)

R R = root, root apex LoS =liquid over solid medium =root

Sc = suspension cells Ls =liquid shake S =shoot

St = internode N = feeder stem, nurse,

- R reservoir medium a =albino

e =embryogenic S =solid medium

v = in vivo SoS = solid over solid medium

? =unknown St = streak plating

S = semi

9 - unknown

tissue culture until few This is due the of a years ago. progress substantially to use

embryogenic cell suspension or seedling tissue, instead of leaf tissue as donor sources for

protoplast isolation.

Species that belong to the Gramineae mostly regenerated into green shoots and

plantlets via embryoid formation. Green corn, fescue, rice and sugarcane plants have

been successfully established in the soil, however, some species regenerated into albino

shoots and plantlets, e.g. Bromus inermis, Festuca arundinacea, Hordeum vulgare, Folium

and Folium and successful perenne multiflorum, Panicum miliaceum Poa pratensis. No

regeneration has been realized for important cereal food crops such as oat, rye and sorghum.

With the which several food also respect to Papilionaceae, to important crops belong,

different species of Glycine (amongst them soybean, G. max), Fotus, Medicago, Pisum and

have shown of the Trifolium recently a regeneration capacity. Leguminous species genera

Arachis, Phaseolus and Vicia, however, are not amenable, so far, for plant regeneration

from protoplasts.

For several woody plant species, representing various families, regeneration procedures have been developed, e.g. Actinidiachinensis (Actinidiaceae), Manihotesculenta (Euphor- biaceae), Firiodendron tulipifera (Magnoliaceae), Populus (Salicaceae), Santalum album

(Santalaceae), Ulmus (Ulmaceae) and different species of the Rosaceae (Malus, Prunus,

Pyrus) and the Rutaceae (

namely Piceaglauca and Pinus taeda, however, many important woody plant species, such as forest trees (Betulaceae, Fagaceae, most of the Gymnospermae and tropical tree species), fruit trees and shrubs (banana, cocoa, coconut, coffee, date, grape and tea) and oil recalcitrant palm, still remain orhave not yet been subjected to protoplast regeneration studies.

PROTOPLAST REGENERATION

Several factors affect the isolation, culture and regeneration from protoplasts, namely the

treatment genotype, the donor tissue and its pre-treatment, the enzyme for protoplast 4 S. ROEST AND L. J. W. GILISSEN

Table 1. (b) Higher plant species (Spermatophyta) that gave development of regenerants from protoplasts of differentdonortissues using several culture techniques

Donor Culture Regenerant

Taxon tissue technique development Reference

CymnospermaeGymnospermae

Pinaceae

Picea glauca eSc B E Attree eletal.al. (1987)

Pinus taedalaeda eSc SS E->PE-*P Gupta & Durzan (1987)(1987)

AngiospermaeAngiospermae Monocotyledonae

Amaryllidaceae

Hemerocallis cv. Sc LsLs C-»P Fitter & Krikorian (1981)

Araceae

Pinellia ternatternal vL Ls C-*E-*PC->E->P Wu etel al. (1986)

Gramineae

Bromus inermis Sc D E-*Pa Kao elal.etal. (1973)(1973) FestucaFesluca arundinacea eSc L C->E-»P,C->E->P, Pa Dalton (1988a,b)

Hordeum vulgare eSc L;S C-*PaC-»Pa LiihrsLiihrs&& L6rz(1988)Lorz (1988)

Lolium eSc L C->Pa Creemers-Molenaar el al. perenne et (1988),

Sc L;B C-»E->PaC->E->Pa Dalton (1988a,(1988a,b)b)

multiflorum eSc L C-*S-»PaC-*S->Pa Dalton (1988b) Oryza sativasaliva Sc S C-*E-»PC->E->P Abdullah etal. (1986),

C LL C-.S-PC->S->P Coulibaly&DemarlyCoulibaly& Demarly (1986),(1986),

Sc L C-PC-*P Fujimura etetal.al. (1985),

Cc BBPP Hayashi elel al. (1986),(1986),

C;Sc B;BN C-»E-*PC-*E-»P Kyozuka etelal.al.((1987),1987),

Sc B;LoS C-»E-»PC-»E->P Thompson etelal.al. (1986),(1986),

Sc L C-.PC-+P Toriyamaelet al. (1986),

Sc L C-S-.PC-.S-.P Yaraada el al. (1986)

Panicum maximum eSc L C-»E-*PC-»E->P LuLu el al. (1981)

miliaceummiiiaceum Sc DD C-E-P,C-.E-P, Pa Heyser(I984)

PennisetumPenniselum americanum eSc D C-*E-*PC-.E-P Vasil & Vasil (1980)

eSc E-.P Vasil al. purpureum eSc D;S E->P Vasil elelal. (1983)

Poa pratensis Sc D C->Sa Van der Valk&Zaal (1988)

Polypogonfugax 7? ?7 C-PC-.P LiLi etal. (1986) Saccharum officinarum Sc B;D;L C-»S-*PC-»S->P Chen elelal.al. (1988)

eSc sS C->E->PC-E-P Srinivasan&Srinivasan & Vasil (1985)

TriticumTrilicum aestivumaeslivum eSc LL C-»E, S-*P Harris elal.etal. (1988)

Zea mays eC LL C-.E-PC-.E-.P Cai etal. (1988),

eSc DN C->E-*PC-E-.P Kamo etal. (1987),

eSc N C->E-»PC-E-P Rhodes eletal.al. (1988),

eSceSc L;S C-EC-.E Vasil & VasilVasil (1987)

Liliaceae

Asparagus officinalis vSt L C-.S Bui-Dang-Ha& Mackenzie (1973)

Haworthia magnifica Sc L C->S-»PC-S->P Sun elet al. (1987) Dicotyledonae

Actinidiaceae

Actinidia chinensischinensis CC L C-S-PC-.S-.P Cai (1988)

Asclepiadaceae Asclepias syriaca Co L C-.S-.PC-S-P Singh (1984) Pergulariapallida Cc L C->S-PC-.S-.P Bapat etelal.at. (1986) Tylophora indica cC L C->E-.PC->E-»P Mhatrecta/.Mhatreelal. (1984) Chenopodiaceae Beta vulgarisvulgaris 7? 7? E-PE->P SteenSteen etetal.al. (1986) Chenopodium glaucum 7? ?7 C-.P Li elet al. (1986)(1986) Compositae

Brachycome dichromosomatica cC L C->S-.PC-S-»P Hahne & Hoffmann (1986)

Chrysanthemummorifolium L N C-»PC-P Otsuka (1986), Otsuka etetal.al.

(1985)(1985) PLANT REGENERATION FROM PROTOPLASTS 5

Table 1. (b) (Continued)

Donor Culture Regenerant

Taxon tissue technique development Reference

CichoriumCichorium endivia L L C->S->PC-S-P Binding el al. (1980,(1980, 1981)

intybus L L C->S->PC-»S-»P Binding elet al. (1981)(1981) Crepis capillaris C L C-»S-»PC->S->P Hahne&Hahne & Hoffmann (1986) DimorphothecaDimorpholheca aurantiaca vL;vSt;Co;C B C-S-»PC->S-*P Al-Atabee & Power (1987, 1988)

Gaillardia grandifloragrandiflora L L C-*S-PC-.S-P Binding etera/.at. (1980, 1981)

L L C-»S->PC-+S-P etal. Helianthus annuusannum L Bindingera/.Binding (1980, 1981)1981)

LactucaLacluca salignasaligna L B C-S-.PC-»S->P Brown elet al. (1987)

sativasaliva vL;L;Co;R L;LoS C-»S-*P Berry el al. (1982),(1982),

vL S C-»S-»PC-S-P Engler & Grogan (1982/1983)(1982/1983)

Petasites japonicus L L C-*S-PC-+S-.P Yabe eletal.al. (1986)(1986) RudbeckiaRudbeckia hirtahirta vL L;B C-»S-»PC-.S-.P Al-Atabee & Power (1987,

1988)

lacinialalaciniata vL;vSt;Co;C B C-*S-PC-S-P Al-Atabee & PowerPower (1988)

B C-*S-PC-.S-P Al-Atabee & PowerPower (1988) purpurea vL;vSt;Co;C SenecioSenecio jacohaea L L C-*S-»PC-S-P Binding elal. (1981) silvalicussilvaticus L L C-S-PC-»S->P Binding etera/.al. (1981)

vernalis L L C->S-PC-S-P BindingBindingsetal.al. (1981, 1982)

viscosus L L C-*S->PC-S-P BindingBinding?/el al. (1981)(1981)

vulgaris L L C-»S->PC-.S-.P Binding & Nehls (1980), Binding?/Binding et al. (1981)

Convolvulaceae

Ipomoea batatas L,StL.St L C->S->PC-S-P Sihachakr & Ducreux (1987)

Cruciferae

ArabidopsisArabidopsis thaliana L L C-*-S->PC-S-P Bindingetel al. (1981),

Sc L C-S-PC-*S-»P Xuan & Menczel (1980)(1980)

BisculellaBiscutella laevigata ?? St C-*S->PC-S-P Bindingelet al. (1988) Brassica alboglabraalboglahra L;Co;R;St L C-*S-»PC-S-P Pua(1987)Pua (1987)

campestriscampeslris Co L C->S->PC-S-P Glimelius (1984)

carinalacarinata Co L C-*S-*PC-S-P Chuong?/etal.al. (1987b)

juncea L L C->E,C—E, S—PS-*P Chatterjeee/o/.Chatterjee?/ al. (1985)

L D C->S-.PC-S-P Kartha elet al. (1974), napusnapus (1974),

vL;L L C->-S-»PC-S-P Thomas eletal.al. (1976),

R LoS C-.S-.PC-S-P XuXu?/el al. (1982b, 1985)

al. nigranigra St LoS C->S->PC-S-P Chuong elet al. (1987a),

Sc L C-EC-*E Klimaszewska & Keller

(1986)(1986)

oleracea Co L C-S-PC-*S->P Vatsya & Bhaskaran (1982),

R LoS C-S-PC-»S->P XuXu?/et al. (1982b, 1985)

Eruca sativasaliva L L C-*E,S-PC—E, S—P SikdarSikdare/etal.al. (1987)

SinapisSinapisalbaalba L L C-*S->PC-S-P Binding et a!.al. (1982)

arvensis L L C->S-*PC-S-P Binding?/etal.at. (1982)

Cucurbitaceae

Cucumus melo L L C->E, S->PS-»P Moreno ?/etal.al. (1986)

al. salivassativus Co B;D C-»E->PC-.E-.P Colijn-Hooymansetet (1988),

L B C-*E-*PC-»E-*P Orczyk & Malepszy (1985)

Euphorbiaceae

ManihotManihol esculenta vL L C-S-PC->S-*P Shahin & ShepardShepard (1980)

Gentianaceae

Gentiana scabra ?? ?7 C->PC-.P Li eletal.al. (1986)(1986)

Geraniaceae

Pelargoniumaridum Sc L C-»S-»PC->S-»P Yarrow et al.at. (1987)

L x hortorumhortorum L L C-»S-.PC-*S-»P Kameya (1975),

Sc L C->S-»PC^S->P Yarrow etel al. (1987)

peltatumpellalum Sc L C-*S-»PC-.S-.P YarrowYarrow?/el al. (1987) 6 S. ROEST AND L. J. W. GILISSEN

Table I. (b) (Continued)

Donor Culture Regenerant Taxon tissue techniquetechnique development Reference

Hypericaceae

Hypericum montbretiimonlbrelii ? St C->S->PC-»S-»P Binding etel at.al. (1988) perforatum ?9 St C-S-»PC-S->P Binding elal.etal. (1988)

Labiatae

MajoranaMajoranahortensis L L C-S->PC-S-*P Binding el at.al. (1982)

Linaceae

Linum alpinum ? St C-*S->PC->S^P Binding etetal.al. (1988), L;CoL;Co B C->E,C->E,S-PS-*P Ling & Binding (1987)(1987) al. amurenseamurense ?7 St C-S-*PC-»S-*P Binding el al. (1988),(1988),

L B C-*S-*PC-S-P Ling & Binding(1987)

hologynumhologynum ?7 St C-»S-»PC-.S-P Binding el al. (1988),(1988), L B C->S-»PC-S->P Ling & Binding(1987)

lewesiilewesii Sc;L L;LoS C->E,C-E.S-PS-»P Barakat&Barakat & Cocking (1985)

?7 St C-»S-»PC-»S->P Binding al. (1988), perenne etel L B C-»S-»PC->S-»P Ling & Binding (1987)

salsoloïdessalsoloides ?7 St C-*S-PC^S-P Binding el al. (1988),

LL B C->S-»PC-»S->P Ling & Binding (1987)

Sc;L L;LoS C S P Barakat&Barakat & strictumstrictum Sc;L L;LoS C-E,S-P-* E, -> Cocking (1985)

usitatissimum Co;R L;LoS C-»S->PC^S-P Barakat&Barakat & Cocking (1983),

L L C->S->PC-S-P BindingsBindingel al.at. (1981,1982),

LL B C->S-»PC-»S-.P Ling & Binding(1987)

Magnoliaceae

Liriodendron lulipiferatulipifera eSc D;N C-»E-PC->E->P Merkle&SommerMerkle & Sommer (1987)

Moraceae

Broussonetia kazinoki L L C-S-PC^S-P Oka &&Ohyama(Ohyama (1985)1985)

Papilionaceae

ClianthusClianlhus formosus LL L C-»S-»PC-»S-*P Binding etel al. (1983)

Crotalaria juncea Co L C->E,C-»E, S-*PS-»P Rao elera/.al. (1982,1985) Dolichos biflorus Sc D C-»EC-.E Sinha&Sinha & Das (1986)

P Glycine canescens Co ?7 P Davey & Power (1988),

Co B C->S->PC-S-P Hammatt el al. (1987),

Co R C-»S-*PC-S-P Newell &&LuuLuu (1985)

clandestinaclandestina Co ? Pp Davey&Davey & Power (1988),

Co B C->S-*PC->S->P Hammatt elelal.al. (1(1987)987)

max Co L C->S->PC->S-P Wei & Xu (1988)

sojasoja Sc D C->EC-E Gamborgc/a/.Gamborg el al. (1983) tabacina Sc D C->EC-E Gamborg elal.«/. (1983) Hedysarum coronarium Co L C-»S-.PC-S-.P Arcioni etel al. (1985b) Lotus corniculatus Co;R L C-»S-»PC-S-P AbujaAhuja el al. (1983a)

tenuis RR L C->S->PC-S-P Piccirilli el a/.al. (1988)

Medicago arboreaarhorea L ?7 C-»S->PC-S-P Arcioni elelal.al. (1985a),

L;R L C-.S-.PC-S^P Mariotti el al. (1984)

coeruleacoerulea L;Sc LoS C->E-»PC—E—P Arcioni etel al. (1982)

difalcatadifalcata Co D;L;N C-E-P Gilmour c/etal.a/. (1987)

falcatafalcala Co D;L;N C—E—P Gilmour elc/ a/.al. (1987)

hemicyclahemicycla Co D;L;N C->E->PC—E—P Gilmour elal.etal. (1987)

glutinosa L;Sc LoS C->E-»PC—E—P Arcioni elelal.al. (1982),

Co D;L;N C-»E->PC—E—P Gilmour etel al. (1987)

salivasativa vL D C->E-»PC-»E-»P Johnson elal.el al. (1981),

vL D E-»PE-P Kao &Michayluk& Michayluk (1980),

Co;R L C-*E->PC-.E-P Lu elal.etal. (1982b),

C;Sc L C-»E-PC-E-.P Mezentsev(l981),Mezentsev (1981),

vL D;LoS C-»E,S-PC-E.S-P Dos Santos elelal.al. (1980),

RR L C->E-»PC-E-.P Xu elal.etal. (1982a)(1982a)

varia Co D;L;N C-»E->PC-*E-»P Gilmour etel al. (1987)

Onobrychis viciifolia vL L;LoS C-*S->P AbujaAhuja elelal.al. (1983b)(1983b)

Pisum sativum vL;Co L;B C-»SC-.S Puonti-Kaerlas & Eriksson (1988) PLANT REGENERATION FROM PROTOPLASTS 7

Table I. (b) (Continued)

Donor Culture Regenerant Taxon tissue technique development Reference

PithecellobiumPilhecellobium dulceduke vL L C->S-*PC-S-P SaxenaSaxena&& Gill (1987)

Psophocarpus tetragonolobusletragonolobus C L C->S-*PC-S-P Wilson el al. (1985), Sc L C->PC-P Zakri (1984)

StylosanthesSlylosatuhes guyanensis Sc L C->S->PC-»S^P Meijer & Steinbiss (1983) Trifolium hybridumhybridum L;R D;L C-+S-.PC->S^P Webb et al. (1984,1986)(1984, 1986)

7? C-.E-.PC-»E-»P & Power pratensepratense Co;L Davey (1988) vL repens vL L;LoS C-»S->PC->S->P Ahuja?/a/.Ahuja et al. (1983b),

Sc D;L C-*S-PC-S-P Gresshoff (1980)

rubens vL;Sc L C-»E-»PC-E-P Grosser & Collins (1984)

Trigonellacorniculata L L C-.E-.PC->E->P Lu el al. (1982a),

L;Sc L C->E-»PC->E->P Dos Santos etelal.al. (1983)

foenum-graecumfoenum-graecum vL D C-SC->S Shekhawat & Galston (1983a)

Vignaaconitifolia vL D C-.E-.PC->E-»P Shekhawat & Galston (1983b)

mungo L D C-.EC->E Sinha?/a/.Sinha et at. (1983)

sinensis sinensis vL L C->E,C->E,SS Davey elet a/,at. (1974)

Ranunculaceae

Nigella arvensis L L C->S->PC-*S->P Bindinget al.al. (1980, 1981)

salivasativa Sc L C-.S-.PC->S-*P Jha& Roy (1982)

Ranunculus sceleratus L ?? C-E-PC-*E->P Dorion & Bigot (1985),

vL L C-E-.P Dorion elc/ al. (1975,1984)(1975, 1984)

ResedaceaeResedaceae

Reseda alba ? St C^S-*PC->S->P Bindingetelal.al. (1988)

lutealulea ? St C-*S-*P Binding elet al.al. (1988)

luteola LL L;St C-*S-*P Binding?////.Bindinget at. (1981,1988)(1981, 1988)

odorata ?7 St C->S->P Binding et al.al. (1988)

phyteuma ?7 St C-.S-PC->S-P Bindinget al.al. (1988)( 1988)

RosaceaeRosacea e

L Fragariaananassa L L C-*S-.PC-*S->P Binding?/Bindingel al.al. (1982)

Malus x domeslica C;Sc L;sS C->EC-»E Kouider?////.Kouider el al. (1984)

Prunus avium x pseudocerasus L;Sc ?? C->R-»PC-»R-*P Davey & Power (1988),

L;Sc Ls C-S-PC-*S->P Ochatt etelal.al. (1987,(1987,1988)1988)

cerasuscerasus L;Sc ? C->R-*PC-»R->P Davey & Power (1988),

L B;L;sS C-R-.PC-*R-»P Ochatt& Power (1988)

7 & Power Pyrus communis L;Sc ? C-*R->PC —> R — P Davey (1988),(1988), vL;L L;sS C->S-*PC-»S->P Ochatt&Caso& Caso (1986)

Rutaceae

Citrus aurantium C sSsS C-»E->-PC-»E-»P Vardi & Spiegel-Roy (1982)

limonUnion C sSsS C-»E-»PC->E-*P Vardi & Spiegel-Roy (1982)

paradisiparodist C sSsS C->E->PC-E-.P Vardi & Spiegel-Roy (1982) reticulatareticulata Cc sSsS C->E-»PC->E-.P Vardi & Spiegel-Roy (1982) sinensis Cc L E-PE-»P Kobayashi etelal.al. (1985), Cc S;sSS;sS C-»E-*P Vardi & Spiegel-Roy (1982),

Vardi etelal.al. (1975)

Vardi?/Vardi elal.al. MicrocitrusMicrocilrus australis x Cc sS C-»E-»PC-»E->P (1986)

australasica

Salicaceae

Populus alba x grandidentatagrandidenlata L L C-»S->PC-S-P Russell & McCown (1986, 1988)

L nigra xx trichocarpa L L C^S^P Russell & McCown (1988) Iremulatremula L LL C-*S-*PC^S^P Russell & McCown (1988)

SantalaceaeSantalaceae

Santalum album Cc LL C-.E-PC->E->P Bapat el al. (1985),

Sc L C-*E-*PC-.E-P Rao & Ozias-Akins (1985)

Scrophulariaceae

Antirrhinum majus vL L C-*EC->E Poirier-Hamon elelal.al. (1974) 8 S. ROEST AND L. J. W. GILISSEN

Table 1. (b) (Continued)

Donor Culture Regenerant Taxon tissue technique development Reference

Digitalis lanata vL L C-»R, S->P Li (1981) obscura L L;LoS C->S-*PC-.S-.P Brisa&Brisa & Segura (1987)

Nemesia strumosa vL L C->S->PC-S->P Hess &Leipoldt& Leipoldt (1979)

Rehmannia gtutinosaglutinosa L LoS C-S-*PC-S-.P Xu & Davey (1983) Solanaceae

Atropabelladonna Sc L C-»E->PC-»E-»P Gosch elal. (1975)

Browallia viscosa Sc LoS C->S->PC-.S-.P Power & Berry (1979)(1979)

Capsicum annuum L D C-*S-*PC-+S-.P Saxena elet al. (1981 b),b),

L B C->S->PC-.S-.P DiazetDiaz el al. (1988)

Datura innoxia L L C->S->PC-.S-.P Schieder(1975,1977)Schieder (1975,1977) metel L L C->S->PC-S-.P Schieder (1977)

meteloïdesmeleloides L L C->S->PC->S-»P Schieder(1977)Schieder (1977)

Hyoscyamus muticus L;Sc L C-.S-.P Ldrzera/.Ldrz etal. (1979),

L L C-*S->PC-.S-.P Wernicke & Thomas (1980),

Wernicke el al. (1979) LycopersiconLycopersicon chilense Sc L C—>S—>PC-S-P Hassanpour-Estahbanati &

Demarly (1986)

esculentumesculenlum Co L C-S-^PC-S-.P Koblitz & Koblitz (1982a,b,(I982a,b,

1983),

C S C-»S->PC-S-.P Morgan& Cocking (1982) peruvianum vL L C-.S-PC-*S^P Muhlbach (1980),

vL L;S C-*E,C-.E.S-PS-*P Zapata & Sink (1981),

Zapata elet al. (1977)

pennellii L;Sc L C-»S-PC-S-P Tan el al. (1987)

pimpinellifoliumpimpinellifolium Co LoS C-*S-»PC-*S->P Imanishi & Suto (1987) Nicotiana acuminata vL LL C-S-.PC->S-»P Bourgin elet al. (1979) alata vL L;S C-»S->PC-S-P Bourgin & Missonier (1978),

Bourgin elal. (1979),

L L C-*S-»PC^s-.P Passiatore&Passiatore & Sink (1981)

honariensis L L C-.S-PC->S-*P Borkird& Sink (1983)

debneyi L S C-»S-.PC-*S-P Piven (1981),(1981),

vL CC-.S-.PP Scowcroft Larkin L;S —* S — & (1980),

vL LL C->S-»PC->S-*P Shakurov(1982)Shakurov (1982)

forgetianaforgeliana L L C-S-»PC-*S-*P Passiatore & Sink (1981)

glauca vL L C-.S-.P Bourgin etelal.al. (1979)

langsdorfii vL L C-»S-»PC-S->P Bourgin el al. (1979),

vL N C-»S-.PC-*S->P Evans (1979)

longiflora vL L C-»S->PC-S-*P Bourgin el al. (1979)

megalosiphon vL L C-.S-.PC->S-P Shakurov (1982)

nesophila vL N C-»S-.PC^S^P Evans (1979)

occidenlalisoccidentalis vL L C->S—>PC->S^P Shakurov (1982)

otophora vL L;S C-*S-*PC-S-.P Banks & Evans (1976),

vL L C->S->PC-S-P Bourgin el al. (1979)

paniculatapaniculala vLvL L C-»S-*PC-S-P Bourgin elal.etal. (1979)

plumbaginifoliaplumbaginifolia vL L C-»S-*PC-S-P Bourgin elet at.al. (1979),

L L C->S-»PC-S-P Gill elelal.al. (1978)

repanda vL N C->S-PC-S-P Evans (1979)

rustica L LoS C-*S->PC-S-P Gill eteial.al. (1979), vL L C-»S->PC-S-P Shakurov (1982) sanderae L L C-*S->PC-S-P Passiatore & Sink (1981)

stocktonii vL N C-»S-»PC-S-P Evans (1979)

suaveolens vL L C-.S-.PC-S-P Bourgin elal.et al. (1979), vL L C-»S->PC-S-P Shakurov (1982)

sylvestris vL L;S C-*S-*PC-S-P Banks & Evans (1976),

vL L C-.S-»PC-S-P Bourgin elelal.al. (1976,1979), L L C-S-»PC-S-P Nagy& Maliga (1976) PLANT REGENERATION FROM PROTOPLASTS 9

Table I. (b) (Continued)

Donor Culture Regenerant

Taxon tissuetissue technique development ReferenceReference

tabacumlabacum vL S C-*S->PC->S->P Nagata & Takebe (1971), Takebe etal. (1971)

velutinavelulina vL L C-*S->PC->S->P Shakurov(1982)Shakurov (1982)

7 ?7 ? Pp Power & Chapman (1983) Nierembergiasp. Petunia alpicola C;Sc L C->S-»PC-S-P Ford-Logan& Sink (1988)

axillaris vL LoS C->S->PC-S-P Power etal. (1976)

hybrida vL;L L C->S->PC-S-P Binding (1974a,b),(I974a.b),

vL L C->S->PC-S-P Durand etal.elal. (1973),

vL S C-*S-»PC-S-P Frearson el al. (1973)

inflalainflata vL LoS C-*S-*PC-S-P Power etal. (1976) parodii vL L;LoS C->S->PC-S-P Hayward& Power (1975) paniflora vL L;LoS;S C-»S-PC-S-P Sink & Power (1977)

violaceae vL LoS C-*S->PC-S-P Power etal. (1976)

Salpiglossis sinuata C L C-»S-*PC-S-P Boyes & Sink (1981), Boyes

elet al. (1980)

Solanum aviculare Sc L C-*S->PC-S-P Gleddie eletal.al. (1985)

brevidens vL L C->S->PC-S-P Barsby & Shepard (1983),

L SoS C-*S-»PC-S-P Haberlach el al. (1985),

L L C->S-»PC-S-P Nelson elal.al. (1983)

cardiophyllum L L;SoS C-»S-»PC-S-P Hunt & Helgeson (personal communication)

chacoense L L;S C-*S-»PC-S-P Butenko &Kuchko& Kuchko (1979)

dulcamara L L C-*S-*PC-S-P Binding & Mordhorst

(1984), Binding& Nehls (1977), Binding el al. (1980,1981)

etuberosum vL LoS C-»S-»PC-S-P Barsby & Shepard (1983),

vL LoS C-»S-*PC-S-P Haberlach etetal.al. (1985)

fernandezianum vL LoS C-»S-*PC-.S-.P Barsby & Shepard (1983)

7 insanum Co ? S-»PS-P Nishio etetal.al. (1987)

khasianum vL L C-»S-»PC-S-P Kowalczyk el al. (1983)

laciniatumtacinialum L;St L C->S-PC-S-P Serraf el al. (1988) luteumluleum L L C-*S->PC-S-P Bindinge/a/.etal. (1980, 1981)

lycopersicoïdeslycopersicoides Sc L C->S-»PC-S-P Handley & Sink (1985),

L;Sc L C-»S-PC-S-P Tan etel al. (1987)

Kumar al. mammosum Sc D;L;sS C-*S-PC-S-P Kumar etel (1983)

melongena vL L C-»S->PC-S-P Bhatt & Fassuliotis(1981),Fassuliotis (1981),

Sc L C-»S-*PC-S-P Gleddie eletal.al. (1982),

L SoS C-»S->PC-.S-.P Guri & Izhar (1984),(1984),

vL L C->S-*PC-S->P Jia & Potrykus (1981),(1981),

L L C-»S->PC-.S-P SaxenaSaxena etal. (1981a, 1987)

nigrumnigrum vL;L L C-*S-PC-.S-P Bindingeletal.al. (1980, 1981), vL;L L C-»S-»PC-.S-»P Nehls (1978)

pennellii L SoS C->S-»PC-»S-P Haberlach el al. (1985), vL L C-»S-*PC-.S-P Hassanpour-Estahbanati& Deraarly (1985)

phureja L;Sc L C-»S-»PC-.S-P Schumann &Koblitz& Koblitz (1983),

Schumann elelal. (1980)

pinnalisectumpinnaliseclum L L C-*S->PC-*S-.P Sidorov elal. (1984)

polyadenium L L;SoS C-»S-»PC-.S-P Hunt & Helgeson (personal communication)

torvum vLvL L;R C->S-*PC-.S-P Guri eletal.al. (1987)

tuberosum L L C-*S-*PC-*S-.P Bindingeletal.al. (1978),

L L C-*S-»PC-.S-.P Bokelmann & Roest (1983),

R D C-»S-*PC-S-P Laine&Laine & Ducreux (1987),

vLvL LoS C-»S->PC-S-P Shepard & Totten (1977)

L D C->S-»PC-.S-P Li & Constabel uporo L D (1984) 10 S. ROEST AND L. J. W. GILISSEN

Table 1. (b) (Continued)

Donor Culture Regenerant

Taxon tissue technique development Reference

verrucosumverrucosum L L C-.S-.PC^S-.P Tan elal.elal. (1987)

xanthocarpum L D C-*S-»PC-.S-+P SaxenaSaxena etelal.a/, (1982)

Ulmaceae

Ulmus x ‘‘Pioneer’Pioneer' CC L C-.S-PC->S-*P SticklenSticklen elal.etal. (1986)

Umbelliferae

Daucus carota ScSc D E-*PE->P Grambow etel al.al. (1972),(1972),

vR Ls C->E-*PC-*E->P KameyaKameya&& Uchimiya (1972)

Foeniculum vulgare ScSc sS E-PE-*P Miura & Tabata (1986)

Levisticum officinale vLvL L C-»E-»P Jia el al. (1988) Ligusticum wallichii Co L C-»E-*PC-E-.P LiLi & Chen (1986)(1986)

isolation, the method ofprotoplast culture, the culture medium, and the physical environment. These factors will be discussed here briefly. For an extensive literature review, the reader is referred to Maheshwari et al. (1986).

For most genotypes ofvarious plant species, regeneration procedures have been developed by trial and error. Therefore, during the last decade, Binding and co-workers have attempted to develop generally applicable protocols. Regeneration capacities ofisolated protoplasts of 77 dicotyledonous plant species were checked under similar experimental conditions (Binding et al. 1981). Cell divisions were obtained in 54 species, giving rise to continuously proliferating callus in 35 species. In 21 ofthese species regeneration ofshoots

found or plants was obtained. From these investigations the following parameters were to be appropriate in various species for successful regeneration: shoot tip regions of aseptic shoot cultures as protoplast sources, high plating densitiesof isolatedprotoplasts, V-KM

and as protoplast culture medium (see the section on ‘The culture medium’) prolonged

of colonies. culture on V-KM agar media, and repeated dilutions the regenerated cell

called ‘streak Recently, Binding el al. (1988) reported on another regeneration procedure plating’. Protoplasts were cultured in streaks of locally high densities. This technique

and of The proved to be advantageous for the culture regeneration many plant species.

of co-workers with factors results Binding and provided more knowledge respect to that play decisive and important roles in the isolation, culture and regeneration ofprotoplasts.

A different successful approach was followed by Cocking and co-workers, who achieved plant regeneration from protoplasts in a variety of plant species by establishing species-specific culture conditions.

Plantfactors

The genotype. Plantregeneration from protoplasts greatly varies from species to species.

In many cases, e.g. tomato, success was limited to a specific genotype under special conditions (Morgan & Cocking 1982). Restoration of the regeneration potential via somatic hybridization in Nicotiana (Maliga et al. 1977) and Lycopersicon (Adams &

Koornneef that the Quiros 1985; et al. 1987; Wijbrandi et al. 1988) suggests regeneration

is dominant which used selectablemarker fusion capacity a trait, can be as a in protoplast experiments.

The donortissue. An increase in the regeneration capacity was observed when protoplasts were isolated from plants that were pre-treated under specific culture conditions (day- length, light intensity, temperatureand relative humidity) in growth chambers (Shepard & PLANT REGENERATION FROM PROTOPLASTS 11

Totten 1977; Kao & Michayluk 1980; Y. Chupeau, personal communication). Binding

(1974b) investigated the effect of the pre-treatment of aseptic shoot cultures of Petunia

hybrida : the yield and plating efficiency of protoplasts were markedly affected by the

composition ofthe medium, the light intensity and temperature during the period ofshoot

culture prior to isolation.

Totipotency is not limited to a certain specialized type of plant cells, since plants have been from derived from regenerated protoplasts all kinds oftissues and organs. Neverthe-

less, the donor tissue used for protoplast isolation has been found to affect greatly the

subsequent culture and regeneration of protoplasts. Meristematic cells from shoot tip

of have been be of regions aseptic grown plantlets proven to a good source protoplasts

with in number of al. a high regeneration capacity a great plant species (Bindings 1981).

In addition, aseptic shootcultures are increasingly used and havebeen demonstratedto be

a good source, e.g. Capsicum annuum (Saxenae/a/. 1981b), Broussonetia kazinoki (Oka &

Ohyama 1985) and Solanum tuberosum (Bokelmann & Roest 1983).

Aseptic shoot cultures and shoot tips are recommended as a source of protoplasts for

several reasons.

(1) The plant material can be easily clonedand prepared which results in a reproducible

high yield of aseptic protoplasts.

The donor tissue is under controlled (2) grown conditions, ensuring a relatively large physiological uniformity and providing fairly consistent results after protoplast

culture.

(3) The protoplasts are isolated from meristematic or incompletely differentiated

tissues, and characterized by an easily inducibleand sustained mitoticactivity and a

high degree of totipotency.

In Dicotyledonae juvenile mesophyll cells of (the apical region of) aseptic shoots, in

in particular, are used as protoplast sources whereas, Monocotyledonae, and woody crops, embryogenic callus or cell suspension cultures, derived from immature embryos, are frequently used. Vasil & Vasil (1980) suggested that the use of embryogenic cultures can overcome the recalcitrance of protoplasts derived from monocotyledons, as demonstrated in Pennisetum americanum. Apart from some mitotic activity, so far no regeneration has been observed in mesophyll protoplasts of monocotyledonous plant species. Protoplasts isolated from embryogenic cell suspension cultures of two woody plant species, both belonging to the Gymnospermae, namely Picea glauca (Attree el al.

1987) and Pinus taeda (Gupta & Durzan 1987), regenerated into somatic embryoids and plantlets. In Picea glauca protoplasts from a non-embryogenic source divided in culture but they never resulted in organogenesis or embryogenesis. It is generally known, however, that during prolonged culture periods suspension cells often lose their morphogenic potential, due to the occurrence of (epi)-genetic instability. As a consequence, new cell suspensions have to be initiated at regular intervals to keep a useful source for isolation of protoplasts with regeneration capacity.

Roots have also be proved to a good source for protoplast isolation. For example, protoplasts isolated from radicles of 1-3 day-old seeds, that were germinated under aseptic conditions, showed a high division potential and callus formationin most of the 12 plant species tested. From these calli, plantlets were regenerated via shoot formationin

Brassica napus, Brassica oleracea and Medicago sativa (Xu et al. 1982a,b, 1985). Plant regeneration was also achieved from root apical protoplasts of Solanumtuberosum (Laine

& Ducreux 1987). Although root protoplasts are generally isolated at relatively low 12 S. ROEST AND L. J. W. GILISSEN

additional which be of quantities, they can provide an experimental system may particular

importance in species from which it is normally difficult to isolate and/or to culture

mesophyll or cell suspension protoplasts.

In species of the Papilionaceae, cotyledonary tissue has been demonstratedto be a good

source for isolation and subsequent regeneration ofprotoplasts.

Externalfactors

The culture medium. An important step in protoplast regeneration appeared to be the

development of the highly enriched 8p culture medium (Kao & Michayluk 1975). The

medium particularly enhanced the development from cultured protoplasts of a meriste-

matic type of cells, which are able to regenerate shoots, as was recently demonstrated

in Capsicum amuum (Diaz et al. 1988). The V-KM medium, composed of the macro elements of the V-47 medium (Binding 1974b) and the other nutrients of the 8p medium,

was developed by Binding and co-workers and proved to be suitable for a variety ofplant

species (Binding et al. 1981), including potato (Bokelmann & Roest 1983).

The physical environment. Chilling freshly isolated mesophyll protoplasts of tomato enhanced the plating efficiency by more than twofold (Miihlbach & Thiele 1981). In

addition, heat shock treatment increased cell division activity of rice protoplasts

(Thompson et al. 1987). Recently, electrostimulationwas also used to enhance protoplast

division and plant regeneration. Electroporation-mediated improvement of cell divisions,

colony formationand regeneration capacity was observed in species ofthe genera Glycine,

Prunus, Pyrus and Solanum (Rech et al. 1987, 1988; Ochatt et al. 1988).

Agarose has been proved to be an important supplement ofthe protoplast culture medium. Apart from avoiding agglutination of protoplasts, plating in agarose stimulates colony formationfromprotoplasts of a wide range of species (Shillito

for et al. 1983). Agarose was also found to be beneficial division of protoplasts of Oryza

al. and wild culture sativa (Thompson et 1986) Medicago species, as compared to in a

medium media solidified with liquid (Gilmour et al. 1987). Using agarose, a stimulating effect was also observed for protoplasts of Primus avium x pseudocerasus (Ochatt et al.

& 1987), Prunus cerasus (Ochatt Power 1988), Pyrus communis and other deciduous woody species (Ochatt & Caso 1986).

embedded in most often cultured in combinationwith Protoplasts agarose are a liquid culture medium. cultured in Liriodendron Protoplasts were agarose droplets, e.g. tulipifera

& Sommer in beads. This latter culture (Merkle 1987), or agarose bead-type technique improved the plating efficiencies in Lycopersicon esculentum and Crepis capillaris, and enabled sustained of of Brassica and Petunia proliferation protoplasts rapa hybrida

(Shillito et al. 1983). This culturetechnique was also successfully employed for protoplast regeneration of Picea glauca (Attree et al. 1987), Cucumis sativus (Orczyk & Malepszy

1985; Colijn-Hooymans et al. 1988), Lactuca saligna (Brown et al. 1987), Dimorphotheca and Rudbeckia(Al-Atabee & Power 1987), Oryza sativa (Hayashi et al. 1986; Kyozuka et al. 1987), Prunus cerasus (Ochatt & Power 1988) and Capsicum annuum (Diaz et al. 1988).

Other techniques employed deal with leafprotoplasts of Populus, which were cultured in contact with polyester screen discs floated in a liquid medium (Russell & McCown

1986). This study represents the first report of reproducible plant regeneration from protoplasts of non-seedling origin of a tree species. Floatationof protoplasts at the air/ PLANT REGENERATION FROM PROTOPLASTS 13

liquid interphase proved to be effective in inducing sustained divisions, callus develop-

ment and ultimate plant regeneration of Brassica carinata (Chuong et al. 1987b). Apart

from the conditioning medium, which was successfully employed for protoplasts of

Liriodendron tulipifera (Merkle & Sommer 1987), feeder-layer techniques have also

proved to be advantageous for the efficient plating of protoplasts at high and low

densities. Galun and co-workers (Raveh et al. 1973) were the first to use X-irradiated

protoplasts of tobacco as feederlayers to support division ofcultured tobacco protoplasts

plated at relatively low densities. Recent examples of the successful use offeeder layers as

nurse cultures for cultured protoplasts concern Petunia hybrida (Shneyour et al. 1984),

differentMedicago species (Gilmour et al. 1987) and Oryza sativa(Kyozuka et al. 1987). In

from addition, somatic hybrid plants were regenerated through the feeder-layer technique

heterokaryons culturedat relatively low densities, which were obtained after electrofusion ofdifferent Solanum species (Puite et al. 1986).

Finally, a microculture technique has been developed, which enabled the culture and regeneration of individual protoplasts of Nicotiana tabacum (Koop & Schweiger 1985) and Brassica napus (Schweiger et al. 1987) in microdroplets in microchambers. This culture technique has been used to develop a sophisticated and automized set-up for

handling single protoplasts (Koop & Spangenberg 1988).

CONCLUSIONS AND PERSPECTIVES

Due to several new approaches concerning plant factors, such as the plant genotype and donor tissue, and external factors such as the culture medium, the physical environment and the appliciation of new culture techniques, considerable progress has been made during the last decade in the field of protoplast regeneration. Until now protoplasts from

212 higher plant species, including several important food crops, belonging to 96 genera of 31 families, have been regenerated to plant(let)s, or in some cases to embryo-like

shoots. these accessible for structures, embryoids or In principle, plant species are genetic manipulation. Plantregeneration has beenreported from protoplasts of several species of the and of which considered be recalci- Gramineae, Papilionaceae woody plants, were to trant. From the data presented in Table 1 and Fig. 1, a further increase in the number of plant species during the forthcoming years is expected. Despite this progress, two major problems still remain and hamper the use of protoplasts for genetic manipulation.

(1) Several economically important crops are not amenable, so far, for plant

regeneration from protoplasts. In addition, in some monocotyledons protoplast

regeneration resulted in the development of albino plantlets only.

(2) A high degree of genetic variation, so-called somaclonal variation, was observed

frequently among the regenerants derived from cultured protoplasts, as reviewed

for potato by Sree Ramulu (1986). Although somaclonal variation might be of

interest for crop improvement (Scowcroft & Larkin 1982), it is undesirable for

genetic manipulation directed towards the additionof desired genetic characters.

Most protoplast regeneration procedures have been developed by an empirical

of far from approach. The diversity protocols clearly demonstrates that we are still under-

the fundamental of standing processes protoplast regeneration. Comparative studies,

like donor culture medium and using experimental parameters tissue, plating density, have been helpful to improve the knowledge of the potentials and requirements ofprotoplasts in culture and have proved to be important for achieving regeneration of many 14 S. ROEST AND L. J. W. GILISSEN

plant species (Binding el al. 1981). However, a real break-through concerning plant

regeneration from protoplasts of recalcitrant species can only be realized when the basic

conditions and activities elucidated. processes leading to organogenic are

Protoplasts of higher plants in culture exhibit a great heterogeneity with respect to the

the characteristics genotype, morphology, physiological and morphogenic capacities.

This heterogeneity of protoplasts in relation to response to culture conditions is a draw-

back in experimental manipulation of higher plant protoplasts (Koop & Spangenberg

1988). Using the microculture technique, developed by Koop & Schweiger (1985), it is

now possible to perform experiments with single protoplasts, the tissue origin and

known. The experimental treatment of which, prior to culture, are precisely individual

tool for funda- culture of protoplasts under controlled conditions represents a powerful

mental studies on the physiology of differentcell types and cell cycle stages, biochemical,

biophysical, cytological and morphological properties, the analysis of differentiation

of al. To programmes and the genetic manipulation protoplasts (Schweiger el 1987).

from of other recalci- realize a break-through concerning plant regeneration protoplasts

fundamental is factors the trant species, more knowledge required on the basic that govern

different in the from to steps regeneration process protoplast plant.

achieved in the fieldof is In spite ofthis fact, the recent progress protoplast regeneration

and allows of via an important step forward genetic manipulation many plant species

such and transfer after fusion several approaches as complete partial genome protoplast

(Negrutiu el al. 1989), direct (Toriyama elal. 1988) and microcell-mediated (Sree Ramulu

al. and & Blaas et 1988) gene transfer, micro-injection (Verhoeven 1988).

ACKNOWLEDGEMENTS

The authors are indebted to Dr Ch. H. Hanisch ten Cate, Dr K. Sree Ramulu, Dr W. J.

Stiekema and Dr L. van Vloten-Doting for valuablesuggestions on the manuscript.

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NOTE ADDED IN PROOF

the of the have been from During preparation manuscript, new data published on plant regeneration protoplasts ofthe following species;

Donor Culture Regenerant

Taxon tissue technique development Reference

Monocotyledonae

Gramineae

Dactylis glomeglomeratarata eSc S C->E-+PC-.E-.P HornetHorn etal.al. (1988)

Triticum aestivumaestivum eC N CC-»E-*Pa-E->Pa Hayashi & ShimamotoShiraamoto

(1988)

Dicotyledonae

Rutaceae

Citrus mitis eSc LoS C->E->PC-*E->P Sim elal.etal. (1988)

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