.'

;I.f;;,,,!.R~;;.93 ilL 96-103 \l990 Pnrrfed in Great Britain

'---~

9b

Ascospore development inCeratocystis moniliformis

P. W. J. VAN WYK

Departmmt of" &rany .md Cmetics. University of the OFS, Bloemfo11trin 9300

:-'1.J. WI1\GFIELD AND P. S. VAN WYK

Depi<rfme71f 0/ .\'-!icrobiotogy and Biocr.£mist;y, Univ£rsrty of th.£ OF5, Bloemlonhin 9300

The uJtrastr cture of perithecium. .1SCUSand ascospore development inCemtocystis 11lOr.lMarmiswas studied and compared with that

or other :\scomycetes. A.scospore fOITnaticn was preceded by the synthesis of delimiting 'n'all structures. associated wirh membranes.These coalescedto form an <lSCUSveside surrounding eight nuclei. The '�eside wan invagiruted to form lobes or sac-like structuresendosing eacn nudeus. Each young ascospore was p.J.ired on one side with a second ascospore, The walIs differenbated to form brimappendages at the tlatter1ed sides of the paired ascosporE'S, Furthermore. the wans de\'elo~d into two layered hat-shaped sheaths. A

third .l!!antoid wail layer as famed towards the inside of the sneath apparently by loma50mes from within the ascospore.i\uxilidr}' cens attached to developing asci and ascospores could bnction in supplying additionai waJl material and nutrients duringspore development. The ultrastructure of C. mOt/lilfarmls is compared with that of CfimbroJra which also has hat-shaped ascospores.

Ce":1tJtYSt;'S SCf!Stllata includes Ophiostoma H. & p, Sydow,

Ceratt'c!,stis Ell. & Haist. Se1lS:1stricto and Caatoc!ShOPSiS

Gpadh. &; Kendr. (De Haag & Scnetter. 198-t: Upadhyay,

1981; Upadhyay & Kenerick. 1975; \Veijman & De Haag,

19;"S). These fungi a:-e adapted to dispersal by insects and

mites (Bndges & \.10ser. 1986; Upadhyay, 1981; von An: &

Van der \-YaJt. 1987; \,vingfie!d & ;...!arasas, 1980).

Light microscope studies of ascocarp developme:1t in

species ofCaafvcystis sa:Sll lato,including the structure or the

perithecium and ascospores, havebee:1 conduded (Andrus.

1930; ,A.ndrus & Harte,. 1933; Baksni. 1951; Dade, 1928;

EliiorL 19:5; Gwynne-\/aughan & Broadhead, 1930: Hutcnin-

son. 1950: \1oreau & \-fareau, 1952; Rosi:1sky. 1961: Sartoris

1927; Tay!or-Vinje. 19,*0,. In contrast. uitrastrudural invesh-

gations have been limited to only a few species. These induce

Opliiostomaste!:oceras (RoDak) Me!in & \.'annf. (Garnson d ai'..

1979), 0. :/{m!' (Buism.; ~bnnf. Ueng & Hubbes. 1980\ and

l- "1111brin:aE!L & Haist. ,Stiers, 1976). Additional cytc!ogica!

studil?5 could tnus contribute to ImprO\"1ng the taxonomy ot

tiliS grouF.

Some 5~ecies ofCer,i:i.'.-ystis SeilS:! :.-;to::.re characterized by

having ascospores with l<.:1usual-shapea sr.eaths. These Include

spores \,vito ineoui1ate:ai. neeale. hat-shaped. or piiiow-

shaped sheaths iUpJdhyay. 19S1). The only other simda!'

a::;cospon;> :or:-:~s .:Ife faura 10 certa!n yeJst genera IBcmconi~.;;1;'. ~9c;"'; Ku:-:zmao &. ,.>,~earn. 197("'. Of the five groups of

,1sca5~'ort;' torms that occur inC:r,'tix~,si'5_ oniy species hilving

~~i:-Shdpt'j sneJtns Ie. ~int1riatalanG t~0S(, \ ith no sheaths:0. ;1:';;iJ.ind 0, s,'mQCir<L~;have been stuJied u!tra~trucruridly.

This study was undertaken to compare the deve!opme!1t and. .shape of ascospores with hat-shaped sheaths in e.nlO'lili/ormis

{Hedgc.! C. Moreau with those of C.{imbnata.

MA TERIAlS A1\D METHODS

An isolate of C. momliformis was obtained from the vl/ood

surface of freshly fe!ledlvlacaranga capensis(BaiH.J Bentn, ex

Sim. NataL South Africa. Cultures were gro\.I,'n on 2°'0 malt

extract agar 120 g Dika y1alt Extract. 20 g DireD Bacto Agar.

I I H.~Oi. at a constant temperature of 18 0e. under uv-light

to promote perimecium formation. Perithecla attached to

small (2 x 4 x 8 a\,'11) blocks of agar, we:-e prepared for

electron microscopy by fixing in 0-1 M (pH 7'0) sodium

phosphate-butTered glutaraldehyde (3 %) for 3 h at 20°.

followed by I h fixation in similarly-buffered OsOI (0'5 '~'o).

The materia! was dehydrated in a graded ethdool series and

embedded in an epoxy resin (Spurr, 19(9). Ultrathin (60 nm)

sections were cut with glass knives. using a LKB !II Ultrotome.

Sections ,yerf' stained for 20 min with uranyl acetate, 10 mm

1;vith lead citrate (Reynolds, 1963'1 and examined with aPhilipsEM300 transmission dectron microscope.

RESUl TS

Electron mic:-ogr2.pnscomprising J. cross section through .:tden~!oping peritnecium showeJ ditte:-ent development.11stages or asci Joel ascospores l:Fig. 1). Asci appeared scatteredthruugflout the ~rithecium. separated by brge dectwn

P.W. ). van Wyk. M. j. Wingfield and P. s. van Wyk 97

Figs 1-3. For capf10n "cc p. 100.

:.,

Ascospore development inC:ratocystis monilifoml/s

Figs ~-,:?; Fur >':J:JtlOnSt:'~p_ IOU

P. W. j. van Wyk. M. j. Wingfield and P. S. van Wyk

~,,"'-:. - ~~,Figs 10-15. for c<:lption set:' F. il1C

Ascospore development inCm1.tvC:Jsfis "1L"1iliformis

transparent areas. These areas contained cell debris. apparentlyarising from degene!"ating mature asci. Closely packed.irregularly.sh.:lped cells. probably ascogenous cells and youngasci. were observed adjacent to the perithecium waU andmature ascospores towards the perithecium centre (Fig. 1).The asa extend from thebase of the perithecium up' to theneck. Asci were not observed on perithecium walls at the baseor the neck (Fig.2'.

Fi e stages in ascospore de elopmentare described. Firstthe young ascus contained a single nucleus. abundantmitocI"!or,l3!"ia and othe!" cellular inclusIOns (Figs 3, 16 AI.Next,

nudeJr divisions resulted in the formation of individual3scospore nuclei (Figs .t 10B). The third stage was defined bythe OCCUITenceof osmiophjJic bodies ~Fiss 5. l6C} and thefourth by the synthesis or an eledron dense spore delimiting

aU 1 Figs O. 16 D).

Delimiting walls of ascospores developed adjacent to andnearly concentric with the ascus wall surrounding most or allcellular inclusions and nuclei (Figs 6, 160). The wallscoalesced and invaginated to enclose each nucleus and othercellula. indusions forming the individual ascospores (Fig. 7). Inlongitudinal section.the~e wallsappeared as sac- or pouch-likestructures sUITounding the nucleus of each developingascospore. The open ends of these sac-like structures wereusually turned towards one side of the ascus before finalascospore delimItation (Figs 7. 16 E). Wall formation wasapparc:ltly associated with membranes (Figs 8, 16 DJ. butassocIation \,,:ith nuclear membranes or endoplasmic retiC'.Jlum,';.10:::-hJtobserved. T o membranes were formed. apparently

rhrou~h de 1!LJ,lOsynthesis, ith an eJectron dense substancebehveen them. The formation of the electron dense substanceappeared to occur simultaneously with d~e production of themembranes themselves (Fig. 9).

1CO

In transve!"se section. eight ascospores could be observed ineach ascus. The ascospores were formed in pairs and adheredto each other at the brim area of the developing wall (Figs 10,16,).

The most prominent featureof the fifth stage of ascosporede-ielopmenl:. was the differentiation of the delimiting wallsinto a distinctive double. layered sheath. enveloping eachspore (Figs 1L 14. 15). Development of the brim of the sheathwas irutiated by the synthesis of thickened highly electrontransparent extensions formed at the edgesand base of thepaired ascospores (Figs I L 16 G)..An inner eledron transparentspore '.vall separated the spore from the sUITounding o::he.1th(Figs H. 15. 16H).

During the fifth stage, unusual auxiliary cells attachedtothe asrus and probab!y the spore wall (Figs 12. 16G~.late=-.these auxiliary cells separated fromthe asci and matureascospores.The area wherethe auxiliaf)' cells were connected

,.to the spore wall was sealed by closureof the epispore tFig.13).A further featureof the fifth developmental stage was the

prese~ceof lomasomes (Figs 14. 15. loG), that appea!"edtocontributeto the formation of the inner wails.After sheathand wall formation was completed, the asci degenerated I,Figs14. IS}and the paired ascospores separated(Fig. IS). This leftindividual mature ascospores amongst the debris of asci andwithin the peritheciaJcavIty. Thesequenceof development ofa~ci and ascospores in C.momhformis is summa!"izeddiagrammatical!y in fig. 16.

DISCliSSIO~

In C. mcnilifomlis young asci appear to be formed fromascogenous cells fo~inS a Hnmg of developing asci adjacentro the perithecium wall. Asci do not appear to be formed in

Figs 1-3. De\'eicpr:;e:-.t.;T:orp:'Oiog:iand u!t.ast;ucture of the?e:ithec-..rr. a:1a asci in C.n:;;r::iifonr.i5 ~'bar= 1 ;.:.r:IJ.Fig. 1. Tra:-'s';e!"se~ection through middle regIon of the pe::thecium, showing cieveio!=,ing, IITepiarly-shaped asci adiacem to the penthecJUm wail a:-.ci

marure asci and aSCO$pores scattered in the centre. fig. 2. Transve:se section. through the neck region of the perithecium. Note lacK of

asci ne.:Jr~he bJse of the neck INB) and sreriJe cushion ceils(Cel. Fig, 3. First developmental stage represented by young ascuscontaimng nude'.Js IN) and several mitochondria(M.L

Figs ~-9. TEy! of developing asci and ascospores in C.mOtlijj[onr.:.sIb.!r = ;)'5 ~m;. fig. 4. ~cond de':eiopmental stage. .-\scus

showing h-,.o or possible eignt nuclei. Fig, 5. Third cieveioprr.ental stage or ascus with oSG"!:ophitic bodies. Fig. 6. Fourth de';elop~entaJ

stage or ascus sr:owing synthesis ofpreiiminary ascospo:o-e dejjrniti~g: wail su.~ounding three n.uclei and cellular inclusions. ~ote..:oa;esdng areas of walls (arTOws.1. Fig. 7. Longitudinal section of .>sc.:.s and Jxospores showing invaginating wails surrounding m.:dei

'.virh sac.like st:"..:ctures tormlng young aseospores. Fig. $.Ot rtOi.'C'synthesis Ct deiimiting \v311 structures. :'-Jote association of rr.e;:,r~ii:;,.inary i'i.111with :;,er::brar.es (a!"raw.'.Fig:. 9. .-\dvanced stage of de:imit:...'"!gwal! and membrane termatien. :-Jete synd-:esis !"eg~ons

.:l!":-OWSjin dose Jssoci.:=.tion with ribosor:-:~siR).

Figs 10-15. TE>! oi de-Hc'iooin!; ascospc:o-€S ,bar = 0'5 :Jml. Fig. 10. T~J.ns';e:-s€ section o~ ~5C''':SJnd :!scospores showing rour;:,a!:s of

~;('unS Jsco~pores JU!"l!1g t:nai stase :111tid~mltatiop.. Fig. 1 1. Pa!r~i. ~.oun~ "!scospores d0~d~ .hs0c:atect dt.:nn~ i0rmation at tr.ecrir::

vi ~!-.e ~h~Jth. Fig. 12. Ste!"ile auxiiiijry cel: whie!' possibiy eomrib1.;:es :,:, Js,:c5?Ore wali fC'f':'-at:on. :-:ote ~uxiHa:y ee1! "ppencia~es

'<1f:"OW'.Fig. 13. .-l,seo~pore sepJrating tram sterile auxiliarycell a=,::enC.:lges. :-\ote 'NJitan:'J le;:'lspOreJ sealing off the area otJ.ppe:1da~..cunnexlun 'J.rrowsJ. Fig. 14. Adherino; pJ.ir('J -Jscospores snowing .je\"e:opn~ent of ~he endospore berween mesospore and

ria~n~.J:~rr:n1.J.Fig. 15. PJireJ ,,~co~pore'~ scparating atter JS(US de~en~J:lOn. :-';ote tne prommt'!1t tI\o.layered sneJth. episporc Ii:'?: J.nJ

,,-t's,)spore ,~...1E;Jnd t!1e endospore ;E:-';~0r spore wail ._LO= !or.~-,soIT':e'.

('"

'f,<pO

~-8°o~B

P. 'vV.J. van Wyk. M. J. Wingfield and P. S. l,rar.Wyk

A

E

101

C)

/ ~"\ ~

~ ,~

\~;r~~\1t:so...pore

Ep!~pore

Endo...port:

Fig. 16. Sche:natic representation of ascus af'\d aKospore deve!opment in C.rtlor.ilifornu5. A. Young ascus wirh dividing nude:..Js(N);

5. :..oung asrus separating from peritheClum waH ;I:h di\'iding nudei; C. iluciear division completed and s:-nthesis of oSr:'joohilic wail

matenal comme:1ces: D_ deiimirjng wan fOrf:1.aho!1 as~C!ated with !T!embrane~; E. ascospore nuciei surrour.cec by sac-iii:.e deJimiting

, ans. oDse;ved in longitudinal section: F. p.m",d a5COS?Ore~ with endosed nuciei; G. steriie auxiJiar:v cell comributing to ascospore

formation; H. degenerating ascus and ascospores ..dhering to each other. L separating IT'_ature ascospore wit~ distinctive :hr~'!ayered

wail

the neck region where the os~jole wiJi develop_ Similarascogenous ceJJs were also reported for O.stmoceras\CaITisonet ai.. 1979), although their specific arrangement was notdiscussed.

A.?rominent feature of ascus and ascospore de\'e!opment inprevIous ultrastmcturai studies of C.5mbriatil (Stiers. 1976),O. stenocenlS1Garrison et aI.. 1979) and 0. :.IImiiJeng &.Hubbes,

1980). is m the mecnanism of spore oeiimitation. In the Jatter

fungi. the ascospore wall is laid down betwee:l a coub!e

delimiting membrane when delimitation commences. Asimj]a; double ascospore delimiting membrane system was

not observed in C Yllonihfom:ls. Ascospore dell..'Tlitation in

C. 71:Ctnii(FC1rnJfsis apparently achieved by wall s~ructures and

:lOr ~embrar.es. Although these independently formed \\,.1]]5

are a5sociated with membranes. a separate double deIimihng

memacanc system. ',\-"<15never obse~.'eci

!t !-:,]5 r.ee:l suggestec ~hat de!imlting memDranes originate

from :he endoplasmic reticulum. nuclear envdope lBt"CKett &

Crawior.i. 1.:170:C:lffoil. 196i'1or tr:e plasmaiemma IBandonid 11.. l>?(7) at t~e aSCL:S.The .ieiimitir.s \\-..;1 sIT'..lCtures in

C. r.:c,;::;{.:m;;=,. ho\';e':cr. appea:- to :,e s:--T.t!1es:::ed.ie r.o~cby a

mec!ta.:lism in\'okmg ribosomes Figs S. -?.\in the ascus(ytO;:-;3S;";1,

Sties ,19~c\ suggested thut ~i\O possib:e methods of

spore delimitation could occur ;:1 C./1mbT1.lt.1""hich. likeC. mom'{ifoTl1-:;s.has hat-shaped ascospores. A double mem-

brane system could fonn a sac a.round eacn nucieus which

enclosed to form individual ascospores. Altematively. the

double memcm,nes formed J large ascus vesideadjace!1t to theascus waJl. This ascus vesIcle the:1 i.r:vaginatec. to torm lobes.

surrounding each nucleus of the young as<:cs!=,ores. We.

however, interpret the formation of an ascus YesicJe around aU

the nudei adjacent to the ascus \\a]L as an early stage in the

delimitation process. The ascus veslde \vouid fic:aHy invaginate

to form sacs which envelop the nucleus o~ each young

ascospore. Depending on the pia:1e in which sections are cut.

the young ascospores wi!! either a??ear e!ong~te and sac-like

in longitudinai section or circular l~ transverse section. Two

separate mec:-:anisms of de!imitaLon JS ?rop0sed by Stiers

(1970i, thus seem improbab:e.

During the 2na! stages ot wa!! ce':e!c::'IT'.e~t::.~eascos?o.-cs

are paired side by side. This pairir.g :s a u:11<:p..:e~eJtu:e or hat-

shaped ascospores and has ,Jjsobee:,. ob5er\"€C 1r'.yeasts s-uci1

as Hm;si'lwi" "il;,~I,lt,ia :HansenJ H &::P. Sy.i. 32"Joni ef ai..

j967. Biack ~ Gorman. 1-?7h ..:.fkr spo,e je!imi[.1:ion.

addltionai '""ait l.1yers are deposlteci be:\';ee~, :he existing

membranes JnJ the wail J.1yer.

Although lomasomes ~re Jp;:,a,endy if'.'.0i\'ed i~l the

:\scospore J('\'eiopmt'nt inCtl'lltoc~Stis rtJol1,[i.fonl1:s

dep0:;,ition of wall material ((ar.-oil. 1967: Sric:-s, 1974;YVilse!1ac~ .$: Kessel. 1965). they were not observed inC iiml>r::f!"u!Stie:-s. 19761. Structures :-esembling lomasomeswero!.howe er, obst'rved in ascospores of C.monili/ormis andit is s:.J~gt'skd that these structures deposit wall material.cont:-ibuting ro inner w;]11(endospore) formation.

The: :i~mber ot wan layers that nave bt'en observed in thedSCO'.ipOrcsot other species ofCml!oc,!shs sa'.$:I!atois variab;e.In C. ,.il'!t'r~nttlthree layers were observed and described as aneiectrvn traospiJrent endospore. a less electron transparent

meSO'5?Cre J:"!d em electron de!lse e?lspore ith an exte:1sion

or brim ,Stiers. 1'?7oi. In C mOlli!i/()nnis the brim of the

Jscospore is an dectmn transparent area. which is apparently

not an extenSIOn or the epispore. The epispore was formed

separJte!y ar.J endosed the e!ectron transparent extension of

the brim. This contrasts with the uniformly electronde!l5e

brim ot C. .nmbno1t«.

In O. wnu with slightly elongated. crescent. shaped

ascos;:,ores .'v.\th no sheaths. 1eng $: Hubbes [1980J described

the ir.:-:e:-electron dense wall as the pnmar: wall and the oute:-

electron t:ar:sp.rent wall as the secondar;... wail. The mesosf'Ore

thus e!ther appears to be absent or was not observed. In

C. mt1mijf~lm!:$ the epi- and mesospore are interpreted as

comtitt:~ing the ascospore sheJth. The endospore would thus

be the spore delimiting wall. Usim;this terminology. it is theas,:ospOie sheath that is hat-shaped and the spore itself thus

has an all.mtoic shaFe.

A. clos~ association appears ~o exist between e!ect:-o~

deme ~oJies (osmiophiJic bod!es,~ a:1d wall formation ir.

e. JI:~,,::i:.:vnmj. Similar structu:es have also ce<=:'r1observed in

S.lcdwfl""j}lfi:' l-crft'151'.liHansen IBecKett et ai.. 1973). O. ;dm:'

IJenS & Hubb:s. l-}SOI and C. .f.mvr:at~ tShers. 197"1.

iIIin~ o:th d II:. (1973) pm\'iced evidence that the lipid

conte:-:.t at the ceils In 5.are~':sil1t. increased ',vith an increase

in b~id ,,'esicles resembling osm!ophilic bodies. Resultsor thisstudy substantiate the suggestion that these bodies function in

'.'.,.all :or7:\ahon.

The h:1al cJI:e:'ermatlon or the ascospore wall or

C. mo/:;;:fLJm:is ,r:~o three distincti':e iaye:-s. 15 cnarac~enze~by the ?resenceot celts connected tothe .asci and ascospore

wails. These auxdiary cells apFear to be similarin snape aCid

size to tne 3S0. :":udear divisior.s and asco:;,pore ceiimihr.g

'..;ail st;""..;,~ures,,\'ere. howen~:-. :lot observed in them. Theythus Jp~ear to be ste:-iJe and possibly playa rolein sheathformJthJn :epi- ar.~ meso spore!. .-\'5 iJ: as we a:-e aware, such

struc::.l:-es han~ nor previousiy bee!1 cDs€:,ved in studies or

aSCO:;,:'0re de\'e:oFlT.ent. The J:.lxdiar:; ceBs couid fur.ction in

pro\'~dir.g JQc!tionai waLl mc1:~:-iJI and r.urrients during

J5Cos;:,ore cevelo;:,mer.t.

Ha~-~n.J~~€-Li J~":osf'.ores Jre :CUi:':: i;::,OC:,:L. ,..;,,:i'r:.~!:,J:1C

C. "!~":!i!f~~MI::5.t;'e two 5rec:es "'!~~: ~he:er.ore hJ\'e r.eer:e.xpec:eJ to na\'e simiiar '.litra:i,r"';Gu;'.1i cn..1rJ..:terisb:s. Hvw-

e\'er. JI:Tl:rer.ces to 'I..J.l!-delimltin~ 5tr\..ictures, aSCQs~orealls. . . "..,

,"'. .'an ::-.t.' 'ie-senc<=:,or -"u.x:!!J:Y (e~!s \\'~a::: :-oune: oenH'en .!':e

~pcc(~:,. This sLl~~es;:s ;:r:J~ 0tr:C 5;:ccics ,it:--. slmi!ar J'5COSF0re

tor[!"":~ :-~ouid h: examlr:I.'J b~:ore (Or..:IUSII r:S can t-e ci~\..r:

fOi +t' g!'Ou~ ..15 J 'snt.llt:'. L'!ti..1Stwccu:'J1 :;,cutiies of

..1S';OS~~0:'oge:ie:,:s 1:1.otht.'r sr-ecle5 0:L't'(,'1:,'l!IStlS snould JJso

102

be underraken. Furtherf!\ore. studiesof these processesinspecies orOor.i(JSrOntllwhich have a more di erse range ofascospore :orms :-ecuires investigabon.

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Cr.lto~r.""l;l~ rimi'~:,ita. (cuma! ..i "\8"T~(1aJ4,,,,iR<,ta"it 46. 10:--:'-1078.

..v.drus. C. r. : 1930\. Ceil relations InL(r;iMstcmd.. I11!Jif.""jlu/i.lt...

.\t;.(,;o:"'~1<126. U3-153.

3.ok5hl. 8. K.( I.;I~ II a~ve]cprne~t or F~r.thecl.:t .l(!J reproouctive ~tructurt'S \fI

~wo specIes orCl""..!rOC:J5115....h:tIlILs vi 8.Jt.",~ 15.33-01.

Bdndoni. R. I.. 8isalputra. A. A. &.: Bisalputra. T. 11907). ,,\scos!"Jfe

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8«j,;e~i:. .0\. &; Craw:urd, R. \.'1. ,19:"Oi_ :\"ude.:tr c~nH'iour and .1~;x-re

ddim:ta:lon ~nX':li.~~';"lfT1' ;:>o.'~"'L'"1';41fOI.ntai or c.rr.~J! .\f:cr<Jv;"Ws:J oj.

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~..:kett. ..\.. Jl[if!~worth. R. F. &: Rose. :\. H. IJ-?7JL Ascospor~ deve1op~t

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se1l.re\;Jtion an.:! ~(!\eio91'1'".E':1t .dur.rI~ .1s.:c>~rorc:s~:1~:m mH.1>~':< ;l ~gt:.

;\rd:lt,tS .of .\I:.-""'v:,':o1:{;' 79. '::J t-l..~.Brid;.;e-s. j, R. &: \Ios~r. I. C. (I.;ISOL Re!iIbonsnip or pr:oretlC m;tes IAQ..'!;

T a~sor.t.'m;dJ~: to the b!u~sta:n run\{us. Crr.lt('t~SI15IIImar ;n rrl"es rnf~ecl~y sou:! m :'If!~ b<;"e-~Je~Coie-.)r1era: St:oiyrlda",.. E'rt'II"i'm>:f"f.li

E'I~''"''A'T.~15, ~~I_.J~j_

CJ:Toil. C. C. .1,)c;). Tite uJtrastrucrure or asco~rore~eiimit,:Hion In ~~,.~

km.'!">u. /":4'1:.1; ," Cril BI<,itJl(!- JJ, .: 1~-'::'::...

Dd~e. H. :\. (1-:11~;.C:;l'al.,):,m:ci!.. ...",!.i,:x,;. the ?ett.'ct stJ~t' oi Ti::(;';"':r:"5~

;".I.,,,j.'I.1 ide~~'n"'5' !'"on

Ho:,nei. :-r".~~,lcr:<JI~ .1' rht B~~,~~h.\b':l"j(~ 3;

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P. 'vV. ]. van Wyk. :Vt J Wingfield and P. S. van YVyk 103

Up.Jdhyay, H P. 1J'?~11. monograph ofCrralocyst:s.wd Cm1forys/iapsl>.

thens: The Univer~lry of Georgia Pfl'~s.Upadhyay. H. P. & Kcndrick. W. B. (]975). Prodromus for a revision of

(enmxvstis!Micro.:JscJies scomycetesl .JndIts conIdia!states..'v1ycofc'SIlJ

117. :"'95-805

\'00 Arx. J. A. & Van Der Walt. J. P (10871. Ophiostol':1ata!es andEndomyceraies 51:dif5 II! ;\1ya,;'oXY30. le7-lib.

Weijman. C. \-1. &: De Hoal;. G. S.r 1-?75'. On the subdivision of the senus

CtrtJU'>CYSliS.,A"iv>I;t' :Jm! Let~wenH.o<,k41. 3S3-JtoJ.

Wi!senach. R. &. Ke~S<:'1.!vi. (10651. The roil" Of ioma~me-s tn waU forrn2tion

in P"I/(i!Iwn; ~'n-»!1(~ll!l;j/I1. io:.mdi of C"'lcriJi .'.1)crci>it1iogy 40. 401-4\)";'

'Sin~field. \1. J. & '\1'-lrilSil5. '\1\'. F. 0. \ 19f.JI. ({TalOC~!!S ips associilted with

Orlhrl~''''')!cJ/s (~(JS!(£ .Coleoptera: Sco!ytidae! on P:r.:.s spp. ifI the CJpe

Province .;:orSm:rf: Africa. PI;'"IDr>nyjadlc.1 12. t5~.

(Received for ~lIbJjmfiol1 16 j\iovel1lber 1989j

I

BRITISHMYCOLOGICALSOCIETY

CENTENARY AWARDS

British Mycological Society support for new initiatives in mycology

The Society's aim is the promotion of the study of fungi. To this end, and to bring to attention the comingcentenary of the Society's founding .in 1896, Council has approved the award of sums of money to individualsand/or groups to support whoJly or in part. activities intended .to promote new initiati..'es in any aspect ofmycology.

.A.ppropriate purposes might be: (i) pilot experiments or observations, especially those covering new ground. tobring a study to a point which might attrad funding from other sources; Iii) support for individuals to aHend formaltraining courses which wilJ enable them to embark on new aspects of their research: (im support for individuals tovisit other laboratories for informal training or collaborative study in a novel topic.

This list is not intended to be exclusive. The emphasis is on support of new initiatives \\'hich, by definition. wecannot foresee. Yau are encouraged to submit an application for an award for such a purpose even jf its intendeduse falls outside the scope implied by the previous paragraph.

Awards wiH not normaily exceed £500. There are no restrictions on eligibility, but these awards ca!U1otbe usedto supplement any postgraduate studentship or bursary. An important condition of the schemeis that awardholders submit a fu!ly receipted accou0t of their use of the moneys within one calendar year of the work beingcompleted and a short report of their mycological activities for possible publication in one of the Society's Journals.

Applications ~UST be made on the form obtainable from the General Secretary. They may be submiHed forconsideration at any time of the year. Applications should be made in advance of any intended expenditure andit would be wise to allow up to six months for the processing of appiications. Application forms can be obtainedfrom Dr A. J. S. Vv'halley.School of Natural Sciences. Liverpool Polytechnic. Byrom Street. Liverpooi 33AF. UnitedKingdom.