Several previously undetected aspects o f the involvement o f oriented fib r e arrays during shape control have emerged from the investigations reported in th is d is s e rta tio n . These are as follow s.
(a) Micronuclear elongation in Paramecium
The overall d is tr ib u tio n o f microtubules during the elongation o f a Paramecium separation spindle has been mapped out fo r the f i r s t time. Elongation involves two types o f microtubules w ith diameters o f about 24 nm and 31 nm. Their d is tr ib u tio n reveals th a t spindle elongation is not due to a simple microtubule s lid in g mechanism.
(b) Ovarian f o l l i c l e shaping in certain insects
Previous studies o f f o l l i c l e development (Tucker & Meats, 1976; Went, 1978) indicated the involvement o f in te rce l 1u la rly aligned microtubules in tension transmission during anisometric f o l l i c l e growth. This study has shown fo r the f i r s t time th a t s im ila r ly aligned microfilaments may also be involved in f o l l i c l e shape control Microfilaments and microtubules can apparently be co-ordinated to produce anisometric f o l l i c l e growth in P. americana. Microfilaments
E x tra c e llu la r matrices have not previously been im plicated in
control o f f o l l i c l e shaping. The f o l l i c u l a r epithelium o f P. americana secretes a f a i r l y substantial e x tra c e llu la r m atrix, the tunica propria. I t consists o f two separate layers - a fibrous layer and a granular layer. The la t t e r has not been detected before. There are
substantial in d ica tion s th a t the tunica propria takes over tension transmission from the e p ith e lia l cytoskeletal system during the la te r v ite llo g e n ic stages o f f o l l i c l e growth in P. americana.
On the basis o f these fin d in g s i t is argued th a t the e x p lo ita tio n o f oriented fib r e arrays may be a widespread feature o f f o l l i c l e
shape control during oogenesis in insects generally. Furthermore, evidence fo r a previously undetected ro le fo r f o l l i c l e c e lls and th e ir cytoskeletons has been obtained. They may enter a 'post- v ite llo g e n ic ' phase in which they a c tiv e ly contract. This seems to form part o f a mechanism fo r discharging mature oocytes from ovarioles a t the s ta r t o f o v ip o s itio n in both o f the insects studied. I t also presumably brings about shape changes associated with corpus luteum formation.
(c) Wing morphogenesis in Calliphora
Waddington's study (1941) o f wing development in Drosophila revealed th a t wing morphogenesis involves a remarkable sequence o f epidermal c e ll shape changes. However, perhaps s u rp ris in g ly , there have been no examinations o f the complete sequence o f fin e stru c tu ra l changes involved in the meantime. Nor have there been investigations o f how each stage in the sequence o f c e ll shape changes is generated.
89 sequence of epidermal c e ll shape changes which is v ir t u a lly
id e n tic a l w ith th a t described fo r Drosophila by Waddington. They also reveal the follo w ing features.
The assembly o f in t e r c e llu la r ly aligned microtubule arrays takes place a t ce rta in points during wing morphogenesis. Their alignments co rrela te w ith d ire ctio n s o f c e ll elongation and/or are related to wing axes. Microfilamentous 'r in g s ' situated at c e ll apices are also involved in shape control at c e rta in stages. Changing patterns o f c e ll ju n ctio n arrangement ind ica te how forces which are generated and/or transm itted by in tr a c e llu la r cytoskeletal arrays may be
d is trib u te d between c e lls during the major c e ll and wing shape
changes. A new proposal is advanced fo r the involvement of filo p o d ia in a 'z ip p e r - lik e ' procedure during the production o f the e p ith e lia l b ila y e r a t the s ta r t o f wing pouch fla tte n in g .
REFERENCES
AGRELL, I.P.S . (1966) C ontinuity o f the membrane systems in the c e lls o f imaginai discs. Z. Z e llfo rsch . mikrosk. Anat. T^, 22-29.
AGRELL, I.P.S. (1968) D iffe re n tia tio n o f the membrane system in c e lls o f imaginai discs. Z. Z e llfo rsch mikrosk. Anat. 365-369.
ANDERSON, E. (1964) Oocyte d iffe r e n tia tio n and vite llo g e n e s is in the roach Periplaneta americana. J. Cell B io l. 131-155.
ANDERSON, L.M. & TELFER, W.H. (1969) A f o l l i c l e c e ll co n trib u tio n to the yolk spheres o f moth oocytes. Tissue and Cell _1_* 533-644.
ASHBURNER, M. & WRIGHT, T. (1978) The Genetics and Biology o f Drosophila. , Vol. 2a. London: Academic Press.
ASHURST, D.E. (1968) The connective tissues o f insects. Ann. Rev. Ent
U : 45-74.
ASHURST, D.E. & CHAPMAN, J.A. (1961) The connective-tissue sheath o f the nervous system o f Locusta m ig ra to ria : an electron microscope study. Quart. J. Micr. Sci. 102: 463-467.
ASHURST, D.E. & COSTIN, N.M. (1974) The development o f a collagenous connective tissue in the locust Locusta m ig ra to ria . Tissue and Cell 6: 279-300.
AUERBACH, C. (1936) The development o f the legs, wings and halteres in wildtype and some mutant stra in s of Drosophila melanogaster. Trans. Roy. Soc. Edinburgh 787-815,
BAKER, P.C. (1965) Fine stru ctu re and morphogenetic movements in the gastrula o f the tree fro g , Hyla r e g i l l a . J. Cell B io l. 95-116
BAKER, P.C. & SCHROEDER, T.E. (1967) Cytoplasmic filam ents and
morphogenic movement in the Amphibian neural tube. Devi B io l. 15: 432-450.
91
BEAULATION, J. (1968) Etude u ltra s tru c tu ra le e t cytochemique des glandes prothoraciques et vers a soie aux quatrième et cinquième âges la rv a ire s . 1 La tunica propria et ses re la tio n s avec les fib re s conjonctives e t les hémocytes. J. U ltra s tru c t. Res. 23: 474-498.
BEHNKE, 0. (1970) Microtubules in disk-shaped blood c e lls . In t. Rev. exp. Path. 1-92.
BEINBRECH, 6. (1970) On the function o f microtubules during the growth periods o f m y o fib rils in insect f l i g h t muscles. Z. Zellforsch. mikrosk. Anat. 109: 138-146.
BELL, W.J. (1969) Continuous and rhythmic reproductive cycle observed in Periplaneta americana. B io l. B u ll, 137: 239-249.
BERNFIELD, M.R., COHN, R.D. & BANERJEC, S.D. (1973) Glycosaminoglycans and e p ith e lia l organ formation. Am. Zool. J_3: 1067-1083.
BERNSTEIN, L.H. & WOLLMAN, S.H. (1976) A circum ferential bundle of microfilaments associated w ith desmosomes near the apex of ty p ic a l th yro id e p ith e lia l c e lls . J. Ultrastruct Res. 326-330.
BODENSTEIN, D. (1950) The post embryonic development o f Drosophila. In Biology o f Drosophila (ed, M. Demerec), pp. 275-367. New York: Wiley and Sons.
BONHAG, P.F. (1959) H istolo g ica l and histochemical studies on the ovary o f the american cockroach Periplaneta americana (L). U. C a lif. Pubis. Ent. J6.: 81-124.
BONHAG, P.F. & ARNOLD, W.J. (1961) Histology, histochem istry and tracheation o f the ovariole sheaths in the american cockroach Periplaneta americana (L). J. Morph. 108: 107-129.
BRINKLEY, B.R. & CARTWRIGHT, J. (1970) Organization o f microtubules in the m ito tic spindle: d if fe r e n t ia l e ffe c ts o f cold shock on microtubule s t a b i li t y . J. Cell B io l. 47: 25a.
BRINKLEY, B.R. & CARTWRIGHT, J. (1975) Cold la b ile and cold stable microtubules in the m ito tic spindle of mammalian c e lls . Ann. N.Y. Acad. Sci. 253: 428-439.
BRYANT, P.J. (1970) Cell lineage relatio n sh ip s in the imaginai wing disc o f Drosophila melanogaster. Devi B io l, 389-411.
BRYANT, P.J. & SCHNEIDERMAN, H.A. (1969) Cell lineage growth and
determination in the imaginai leg discs o f Drosophila melanogaster. Devi B io l. 263-290.
BUNGE, M.B. (1973) The fin e stru ctu re o f nerve fib re s and growth cones o f isolated sympathetic neurons in cu ltu re . J. Cell B io l. 56: 713-735.
BURGESS, D.R. & SCHROEDER, T.E. (1979) The cytoskeleton and cyto- musculature in embryogenesis - an overview. Meth. Achiev. exp.
Pathol. 8: 171-189.
BURNSIDE, B. (1971) Microtubules and microfilaments in newt neurulation Devi B io l. 416-441.
BURNSIDE, B. (1973) Microtubules and microfilaments in amphibian neurulation. Am. Zool. 1_3: 989-1006,
BUXTON, P.A. (1930) Resemblance between a pholcid spider, a t ip u lid , and a reduviid in Samoa. Proc. Ent. Soc. Lond. 2r. 6566.
BYERS, H.R., FUJIWARA, K. & PORTER, K.R. (1980) V isu a liza tio n o f microtubules of c e lls in s itu by in d ire c t immunoflorescence. Proc. Natl. Acad. Sci. (U.S.A.) 77: 6657-6661.
BYERS, B. & PORTER, K.R. (1964) Oriented microtubules in elongating c e lls o f the developing lens rudiment a fte r induction. Proc. Natl. Acad. Sci. U.S.A. 1091-1099.
CANDE, W.Z. (1981) Physiology o f chromosome movement in lysed ce ll models. In In te rn a tio na l Cell Biology 1980-81. (ed. V.G. Schweiger) pp. 382-391, B e rlin : Springer-Veriag.
CANDE, W.Z. (1982) Nucleotide requirements fo r Anaphase chromosome movements in permeabi1ized m ito tic c e lls : Anaphase B but not Anaphase A requires ATP. Cell 28: 15-22.
93
CHANG, M.C. & GOLDMAN, R.D. (1973) The lo c a liz a tio n o f a c tin - lik e fib e rs in cultured neuroblastoma c e lls as revealed by HMM binding. J. Cell B io l. 867-874.
CHAPMAN, R.F. (1972) The Insects : Structure and Function. (ed. W.S. Bui lough) London: English Univ. Press.
CLARKE, M. & SPUDICH, J.A. (1977) Non muscle c o n tra c tile proteins: The role of a ctin and myosin in c e ll m o tility and shape determina tio n . Ann. Rev. Biochem. 797-822.
CLONEY, R.A. (1966) Cytoplasmic filam ents and c e ll movements: Epidermal c e lls during ascidian metamorphosis. J. U ltra s tru c t. Res. T4: 300-328.
CLONEY, R.A. (1972) Cytoplasmic filam ents and morphogenesis : e ffe cts o f cytochalasin B in c o n tra c tile epidermal c e lls . Z. Z e llfo rsch . Mikrosk. Anat. 1 ^ : 167-192.
COHEN, A.M. & HAY, E.D. (1971) Secretion of collagen by embryonic neuroepithelium at the time o f spinal c ^ rd -s o m ite in te ra c tio n . Devi B io l. 578-605,
COHEN, J ., BEISSON, J. & TUCKER, J.B. (1980) Abnormal microtubule de ployment during defective macronuclear d iv is io n in a Parameciurn mutant. J . Cell Sci. 153-167.
CONKLIN, E.G. (1932) The embryology o f Amphioxus. J, Morph. 54: 69-118,
COOKE, P. (1976) A filamentous cytoskeleton in vertebrate smooth muscle fib e rs , J. Cell B io l. 68: 539-556.
CROSSLEY, A.C.S. (1964) An experimental analysis o f the o rig in s and physiology o f haemocytes in the blue blow fly Cal1iphora
erythrocephala. J. exp. Zool. 157: 375-398,
DAVEY, K.G. (1965) Reproduction in the in se c ts . London: O liver & Boyd Ltd.
DAVIDSON, L. & LAFOUNTAIN, J.R. (1976) Preliminary observations on the u ltra s tru c tu re o f the macronuclear a m ito tic and micronuclear m ito tic and m eiotic spindle apparatuses in the p rim itiv e eukaryote Tetrahymena-pyriformis. J. Protozoology 8A.
DE BIASI, S. & PILOTTO, F. (1976) U ltra s tru c tu ra l study o f collagenous structures in some Diptera. J. Submicr. C yto l. 8: 337-345.
DE WILDE, J. & DE LOOF, A. (1974) Reproduction. In The Physiology of Insecta. , 2nd e d itio n . (ed. M. Rockstein) New York: Academic Press
DIPPELL, R.V. (1955) A temporary sta in fo r Paramecium and other protozoa. Stain Technol. 60-71.
DUNLAP-PIANKA, H.L. (1979) Ovarian dynamics in Heliconius b u tte r flie s : co rrela tio ns among d a ily o v ip o s itio n rates, egg weights and quanti ta tiv e aspects o f oogenesis. J. Insect Physiol. 741-749.
DUNN, G.A. & HEATH, O.P. (1976) A new hypothesis o f contact-guidance in tissue c e lls . Exp. Cell Res. 101: 1-14.
DUSTIN, P. (1978) M icrotubules. New York: Springer-Verlag.
EDELMAN, G.M. (1977) Transmembrane control and surface modulation in animal c e lls . Prog. C lin . B io l, Res, VI_\ 467-480.
EDWARDS, J.G. & DYSART, J. McK, (1980) M u ltic e llu la r cytoskeletons o f BHK 21 c e lls . Cell B io l. I n t n l . Rpts. 4: 729.
EDWARDS, U.S., MILNER, M.J. & CHEN, S.W. (1978) Integument and
sensory nerve d iffe r e n tia tio n o f Drosophila leg and wing imaginai discs in v i t r o . Wilhelm Roux's Arch. Devi B io l. 185: 59-77.
FAUSTO-STIRLING, A. (1978) Pattern formation in the wing veins o f the fused mutant (Drosophila melanogaster). Devi B io l. 63: 358-369.
FAWCETT, D.W. (1966) An a tla s o f fin e stru ctu re . The C ell. Its organelles and inclu sion s. London: W.B. Saunders Co.
95
FEKETE, E., FRISTROM, D ,, KISS, I. & FRISTROM, J.W. (1975) The
mechanism o f évagination o f imaginai discs o f Drosophila melanogaster. I I Studies on trypsin-accelerated évagination. Wilhelm Roux's
Arch. Devi B io l. 178: 123-138.
FISCHMAN, D.A. (1967) An electron microscope study o f m yo fib ril formation in embryonic chick skeletal muscle. J. Cell B io l. 32: 557.
FORER, A. (1976) Actin filam ents and b iré frin g e n t spindle fib re s during chromosome movements. Cold Spring Harbor Conf. Cell P ro life ra tio n 3C: 1273-1293.
FRISTROM, D. (1976) The mechanisms o f évagination o f imaginai discs o f Drosophila melanogaster. I I I . Evidence fo r c e ll rearrangement. Devi B io l. M : 163-171,
FRISTROM, J.W. (1972) The biochemistry o f imaginai disk development. In The Biology o f Imaginai Disks., Vol. 5. (eds. H. Ursprung & R. Nothiger) pp, 109-154. B e rlin : Springer-Verlag.
FRISTROM, D.K., FEKETE, E, & FRISTROM, J,W. (1981) Imaginai disc development in a non-pupariating leth al mutant in Drosophila melanogaster. Wilhelm Roux's Arch. Devi B io l. 190: 11-21.
FRISTROM, D. & FRISTROM, J.W, (1975) The mechanism o f évagination o f imaginai discs o f Drosophila melanogaster. 1. General considerations Devi B io l. 4^: 1-23,
FUJITA, S. (1960) M ito tic pattern and histogenesis o f the central nervous system. Nature (Lon.) 185: 702-3.
GARCIA-BELLIDO, A. (1975) In Cell P a tterning. Ciba Foundation Symp. 161-182.
GARCIA-BELLIDO, A. (1977) Inductive mechanisms in the process o f wing vein formation. Wilhelm Roux's Arch. Devi B io l. 182: 93-106.
GARDINER, B.G.C. & MADDRELL, S.H.P. (1972) Techniques fo r routine and large scale rearing of Rhodnius prolixus Stal (Hemiptera Reduviidae). B u ll. ent. Res. 505-515.
GIBBINS, J.R ., TILNEY, L.G. & PORTER, K.R. (1969) Microtubules in ,th e formation and development o f the primary mesenchyme in Arbacia punctata. I The d is tr ib u tio n o f microtubules. J. Cell B io l. 41 : 201-206.
GOLDMAN, R.D,, POLLARD, T.D. & ROSENBAUM, J. (1976) Cell M o t i li t y . New York: Cold Spring Harbor,
GRANT, P. (1978) Biology o f Developing Systems. New York: H olt, , Reinhart & Winston.
GUSTAFSON, T. & WOLPERT, L. (1967) C e llu la r movement and contact in sea urchin morphogenesis. B io l. Rev. 442-498.
GUTHRIE, D.M. & TINDALL, A.R. (1968) The Biology o f the Cockroach. London: Edward Arnold,
HAUSER, M. (1972) The in tra n u cle a r m itosis o f the c ilia te s Paracineta 1imbata and Ich th yo p h tiriu s m u l t i f i l i i s . I, Electron microscope observations on pre-metaphase stages, Chromosoma 158-175.
HAY, E.D. (1973) Origin and ro le o f collagen in the embryo. Am. Zool. 13: 1085-1107.
HEATH, I.B . (1980) Variant mitoses in lower eukaryotes - indicators o f the evolution o f m ito sis, In t. Rev. Cytol. 64: 1-80.
HEUBNER, E, & ANDERSON, E, (1970) The e ffe cts of v in b la s tin e su lfa te on the m icrotubular organisation of the ovary o f Rhodnius p ro lix u s . J. Cell B io l. 191-198.
HEUBNER, E. & ANDERSON, E. (1972 a) A cyto lo gica l study o f the ovary o f Rhodnius p ro lix u s . Part I.The ontogeny o f the f o l l i c u l a r
97
HEUBNER, E. & ANDERSON, E. (1972 b) A cytological study o f the ovary o f Rhodnius p ro lix u s . Part II. Oocyte d if fe r e n tia tio n . J. Morph.