10
PREVIOUS WORK
The previous investigations on the embryology o f
t h i s family h a v e been s u m m a r i s e d by Schnarf(l929,1931),
Davis(1966),
~ h a h ( 1 9 7 2 ) and N i j a l i n g a p p a a n d
Nagaraj
(19801,
The e m b r y o l o g i c a l work o n C y p e r a c e a e d a t e s b a c k t o
1 8 1 0 when Mir.bel (cited in B r a u n 1 8 6 0 ) r e p o r t e d a n u n u s u a l
o c c u r r e n c e o f t h i n e m b r y o s i n C a r e x p e n d u l a , Ther.eafter,
for nearly f i v e d e c a d e s , hardly
any progress was made
until 1 8 5 8 when H o f m e i s t e r d e s c r i b e d t h e e m b r y o s a c s o f
C. arenaria.
-
C, hirta
a n d C. p a n i c e a , I n 1 8 6 1 , h e m a d e
..
some observations on the embryo development in
Vescue(1879),
C. p a n i c e a .
F i s c h e r .(188O) and J o n s s o n ( l 8 8 1 ) r e p o r t e d
-
t h e "Normal" t y p e o f e m b r y o s a c s i n C . v e s i c a r i e , C. p r a e c o x
and C. a c u t a r e s p e c t i v e l y a n d c o n f i r m e d H o f m e i s t e r ' s fin-
.
d i n g s in p t h e r s p e c i e s o f Cdrex. W h i l e t h e s e p i o n e e r i n g
workers concentrated their studieson the female reproductive
s t r u c t u r e , Elfving(1879) paid a t t e n t i o n t o t h e m a l e r e p r o d u c t i v e structure' and
described the development o f
p s e u d o m o n e d i n s o m e species, o f C a r e x , C y p e r u s a n d E l e o c h a r i s
This mode o f development o f the male gametophyte turned
out t o be quite unique among the angiosperms and
/
prompted Wille(1882, 1 8 8 6 ) a n d S t r a s b u r g e r ( l 8 8 4 ) t o
r e i n v e s t i g a t e &I
palustria and confirm certain obser-
v a t i o n s m a d e e a r l i e r by ElFving. W h i l e E l f v i n g a n d
Straaburger considered that the abortive nuclei become
r e e o r b e d 'by t h e c y t o p l a s m o f t h e t e t r a d , W i l l e assumed t h e i r
f u s i o n with t h e f u n c t i o n a l nucleus.
A f t e r a few y e a r s ,
Wilczek(1892) described t h e
structure
o f embryo . i n a n u n d e r t e r m i n e d s p e c i e s o f S c i r p u s a n d s k e t c h e d
some s t a g e s o f d e v e l o p m e n t b u t e r r o n e o u s l y i n t e r p r e t e d t h e
c o t y l e d o n a r y s h e a t h a s t h e r a d i c l e and t h e t r u e r a d i c l e a s
an a d d i t i o n a l r o o t . However, h e gave t h e c o r r e c t i n t e r p r e t a t i o n i n r e s p e c t o f t h e p l u m u l a r p o r t i o n o f t h e embryo b y
d i f f e r e n t i a t i n g t h e f i r s t l e a f from t h e e p i c o t ' y l a r y m e r i s t e m .
Then f o l l o w s a more s y s t e m a t i c and p a i n s t a k i n g s t u d y on
and g e r m i n a t i o n b y D i d r i c h s e n ( 1894, 1 8 9 7 ) .
embryo development
He s t u d i e d a s many a s 2 8 s p e c i e s o f 1 6 g e n e r a and
rectified
an e r r o r i n i n t e r p r e t a t i o n o f c o t y l e d o n a r y s h e a t h end r a d i c l e
i n S c i r p u s embryo made b y W i l c z e k i n 1 8 9 2 . He was t h e f i r s t
,
.
t o g i v e t h e c l a s s i f i c a t i o n of
the basis o f disposition o f
embryo i n C y p e r a c e a e on
d i f f e r e n t embryonal organs
i n t h e m a t u r e embryo.He r e f e r r e d t h e embryo t y p e s e d n i n
Carex v u l p i n a t o t h e Cerex
-
t y p e and t h a t s e e n i n C y p e r u s
m u c r o n a t u s t o t h e Cyperus t y p e ,
the other.taxa studied
c o r r e s p o n d i n g t o e i t h e r o f t h e s e two t y p e s ,
With t h e s t a r t o f t h e present century,
several authors
w i t h a c y t o l o g i c a l b e n t o f m i n d began t o a n a l y s e
t h e course
of p o l l e n development i n s e v e r a l s p e c i e s o f Cyperaceae
from a c y t o l o g i c a l a n g l e . T h e y f o c u s s e d t h e i r a t t e n t i o n
on t h e mode o f c y t o k i n e s i s i n t h e m i c r o s p o r e m o t h e r c e l l ,
t h e n a t u r e QF s e p t a s e p a r a t i n g t h e m i c r o s p o r e n u c l e i i n a
tetrad, t h e f o r m a t i o n o f g e n e r a t i v e c e l l a n d .sperm c e l l s and
degeneration o f abortive microspores during the formation o f
pseudomonad. ~ u e l ( 1 9 0 0 ) s t u d y i n g t h e m i c r o s p o r o g e n e s i s a n d
male g a m e t o p h y t e i n C a r e x a c u t a r e p o r t e d t h e f o r m a t i o n o f
an e p h e r m e r a l c e l l p l a t e d u r i n g i n t e r k i n e s i s i n m i c r o s p o r e
mother c e l l a n d c o m p l e t i o n o f first p o l l e n m i t o s i s e v e n i n
t h e a b o r t i v e microspores. S u b s e q u e n t l y , s i m i l a r s t u d i e s w e r e
extended to the other species o f Carex, Cyperus, Eleocharis,
Fuirena, Isolepis and Scirpus (Stout 1912; Heilborn 1918;
S u e s s e n g u t h , 1919; P i e c h 1924a, 1 9 2 4 b , 1 9 2 8 ; ~ a k a n s s o n 1 9 2 8 ,
1951; W u l f f 1 9 3 9 ; M a r t e n s 1 9 3 9
a n d P f e i f f e r 1942)..Heilborn
( 1 9 1 8 ) and Suessenguth(l919), i n a d d i t i o n t o t h e p o l l e n
development, h a v e s t u d i e d t h e f o r m a t i o n o f e m b r y o s a c a n d
e n d o s p e r m a n d they were t h e f i r s t t o r k p o r t t h e e n d o s p e r m
d e v e l o p m e n t i n Cyperaceae. Martens(q9391, h o w e v e r , d e s c r i b e d
the development o f male gametophyte and embryo sac in Carex
picta.
..
The i n v e s t i g a t i o n s c o n c e r n i n g e m b r y o g e n y o f C y p e r a c e a e
were r e v i v e d by S c h n e i d e r ( 1 9 3 2 ) w h o m a d e a d e t a i l e d s t u d y
of the development o f embryo and germination in eight
s p e c i e s b e l o n g i n g t o G r e x , C y p e r u s , Fimbristylis., F u i r e n a ,
Kyllinaa, S c i r p u s a n d Scleria. S h e c l a s s i f i e d t h e m a t u r e
embryos seen in these taxa into the Carex type, t h e Cyperus
type and the Scirpus type o f which the first two were
p r o p o s e d by Didrichsen(l897).
Another major contribution
during this period i s that o f Tanaka (1939-1941) who made
a n e x t e n s i v e s t u d y o f t h e p o l l e n development i n severa'l,
Japanese s p e c i e s o f B u l b o s t y l i s , -9Carex
Rhynchospora,
Cyperus,
Kvllinqa
S c i r p u s and S c l e r i a . L a t e r , h e ( 1 9 5 0 ) d e s c r i b e d
t h e d e v e l o p m e n t o f embryo s a c i n t h r e e s p e c i e s o f C a r e x .
U n l i k e t h e p r e v i o u s a u t h o r s who a p p r o a c h e d t h e p r o b l e m s
w i t h a c l a s s i c a l p e r s p e c t i v e and b y c o n f i n i n g o n l y t o one
o r t w o a s p e c t s o f s t u d y , many w o r k e r s a f t e r 1950 p o s s i b l y
i n s p i r e d b y t h e works o f S c h n a r f ( 1 9 2 9 ,
1 9 3 1 ) and Maheshwari
( 1 9 5 0 ) h a v e made e a r n e s t e f f o r t s t o g a t h e r a s much i n f o r m a t i o n
as p o s s i b l e w i t h a v i e w t o c o m p i l e a l l e m b r y o l o g i c a l f e a t u r e s
o f a g i v e n t a x o n and t o a p p l y them f o r t a x o n o m i c p u r p o s e s .
Dnyansagar and T i w a r i ( l 9 5 6 a ,
1956b) were t h e f i r s t
t o i n i t i a t e e m b r y o l o g i c a l s t u d i e s o f Cyperaceae i n I n d i a .
They d e s c r i b e d t h e o n t o g e n y o f a n t h e r w a l l ,
m i c r o - and
m e g a s p o r o g e n e s i s and t h e d e v e l o p m e n t o f m a l e and f e m a l e
gametophytes i n F i m b r i s t y l i s
quinquanqularis.
and S e m l o w ( l 9 5 7 ) , on t h e o t h e r hand,
Guttenberg
t a b k l e d t h e problem
r e l a t e d t o ontogeny a n d . h i s t o g e n i c p a t t e r n i n plumule,
r a d i c l e and m e s o c o t y l in.Carex
Later,
a r e n a r i a and C y p e r u s f e r o x .
Padhye and h i s a s s o c i a t e s s t u d i e d t h e e m b r y o l o g y
o f e i g h t s p e c i e s b e l o n g i n g t o Cyperus, E l e o c h a r i s ,
Fimbristylis,
K y l l i n g a and ~ c i r p u send d i s c u ' s s e d t h e
a f f i n i t i e s between Cyperaceae and Junacaceae i n t h e l i g h t
o f e m b r y o l o g i c a l s i m i l a r i t i e s ( Padhye 1959,
1971a, 1971b; Padhye and M o h a r i r 1958
1960,
1968,
end Padhye,Sandhya
a n d A n n a p u r n i 1 9 7 0 ) . D u r i n g t h e same p e r i o d ,
Shah and h i s
c o - w o r k e r s t o o k u p s i m i l a r i n v e s t i g a t i o n s b y p a y i n g more
attention t o microsporogenesis, pollen development,embryogeny
and seed coat and pericarp. They studied eight s p e c i e s
belonging t o seven genera, Bulbostylis, Carex, Cyperus,
Eleocharis, Kyllinsa,
Scirpus and Scleria (Shah1959, 19628,
1962b, 1962c, 1964, 1965, 1967 , ' 1 9 6 8 ; P a t e 1
a
and Shah 1 9 6 0 ,
1962; Shah and Neelakandan 1971). Shah(1965) suggested
a new embryo t y p e t o accommodate t h e e m b r y o observed i n
Bulbostylis barbata and considered t h e C y p e r u s and Scirpus
types o f Schneider ( 1 9 3 2 ) as identical and t r e a t e d the
Scirpus type a s invalid. Guignard ( 1 9 6 1 ) studied t h e embryogeny of Cyperus vegetus
and
traced the d e r i v a t i o n s of
different embryonal regions by following s u c c e s s i v e s t a g e s
o f embryo de,velopment,Khanna(1963, 1965) investigated
embryology o f Cyperus rotundus,
the
C. triceps, Kyllinqa
melanosperma and Scirpus mucronatus and for t h e f i r s t t i m e
in Cyperaceee, reported the Asterad t y p e o f e m b r y o development
in
mucronatus which on reinvestigation w a s s h o w n t o be
incorrect' ( Nijalingappa and Tejavathi 1977). Another notable .
contr'ibution on pollen development w a s m a d e by C a r n i e 1 ( 1 962).
He presented a lucid account o f t h e development o f anther
wall and male gametophyte in Carex hirta, C ~ e r u sa l t e r n i f o l i u s
Eleocharis marnillata, Kyllinaa t r i c e p s and S c h o e n o p l e c t u s
tabernaemontani and on t h e b a s i s o f c e r t a i n s i m i l a r i t i e s
in respect o f microsporogenesis i n C y p e r a c e a e and Juncaceae,
concluded that t h e family Cyperaceae i s closely r e l a t e d .to
the Junacaceae than t o the Gramineae. Grezicka ( 1 9 6 4 ) reported
the megasporogenesis and the development o f embryo sac in
Cardx aristata.
-
The most exhaustive embryological account t h u s
far known in the present century is that o f V,an der Veken
1965). H e examined t h e mature embryos o f 3 4 2 s p e c i e s o f
Cyperoideae and found six main types which were named t h e
Cyperus, Carex, Schoenus, Fimbristylis, B u l b o s t y l i s and
Scirpus types; further, embryographic data were used t o split
certain heterogeneous genera such a s Scirpus and Fimbristylis
into
several smaller genera .Subsequently, s i m i l a r s t u d i e s
were extended t o t h e species o f
~h~nchosporineae(Verbe1en
1970), Cladiinae, Gahniinae ( , V a n h e c k e 1974) and Bisboeckelereae
(Meert and Goetghebeur 1979). Concomitantly, Strandhede
'
(1965a, 1965b, 1 9 6 5 ~ )during h i s extensive cytotaxonomic
survey o f some European species o f Eleocharis reported t h e
development o f male gametophyte in
and
E.
mamillata,
E.
palustris
E. uniulumis. On the contrary, Juguet end h i s c o w o r k e r s
gave a detailed account o f embryo development i n several
cyperaceous taxa and used their findings for t a x o n o m i c
purposes (Juguet 1966a, 1966b, 1967, 1 9 6 9 1970a, 1970b,
1971, 1972, 1973; Juguet end L e b e g u e 1966 a n d Juguet and
Vallade 1979). Another serieb o f papers h a v e m a p p e a r e d o n
t h e embryology o f Indian Cyperaceae by N a g a r a j a n d his
pupils.They
examined 1 8 species with particular a t t e n t i o n
to s o m e genera s u c h a s Ascopholis, Diplacrum, Fimbristylis,
Fuirena, Hy~olytrurn, Lipocarphs, ~ h y n c h 0 8 p o r a , ~ c i r p u sa n d
~ c l a r i a . whoar embryology has been unknown o r incompletely know1
(Nagaraj and Nijalingappa 1968, 1972, 1973; Nagaraj
Nijalingappa and Tejavathi 1976; Nijalingappa 1976, 1977c,
1986; Nijalingappa and Tejavathi 1977, 1983; Nijalingappa
and Devaki 1978, 1979; Nijalingappa and A r e k a l l 9 8 3 ; Tejavathi
1907). In these investigations, attempts have been made t o
critically e x a m i n e the, topographic relations and t h e o r i g i n
o f t h e epicotyl & t h e
cotyledon in t h e e m b r y o o n the l i n e s
o f Haccius(l952) and Swamy ( 1 9 6 3 ) and established that both
the epicotyl and t h e cotyledon arise from t w o adjacent
sectors o f the terminal tier o f the proembryo a s r e p o r t e d
for several other monocotyledonous taxa. In addi.tion, Nijalingappa and Devaki ( 1 9 7 8 ) reported t h e e n d o s p e r m haustoria
'in Scleria folibsa for t h e first t i m e in t h e family
Cyperaceae.
All embryological investigations done u p . t o 1 9 7 0 were
based entirely o n light microscopic o b s e r v a t i o n s and a
few o f t h e p a p e r s which appeared t h e r e a f t e r dealt with
electron microscopic studies. Carnie1 (1971a,197.lb) was
the first t o undertake ultrastructural s t u d i e s o f C y p e r a c e a e
using E l e o c h a r i s palustris, t h e most f a v o u r i t e m a t e r i a l
used for light microscopic i n v e s t i g a t i o n s by e a r l i e r
workers(E1fving 1 8 7 9 ; Strasburger 1 8 8 4 ; S t r a n d h e d e 1965a,
and Thiebaud 1970). H e (1971a, 1971b) s t u d i e d t h e f o r m a t i o n
o f pollenkitt a n d z b i s c h bodies and t r a c e d a l l e v e n t s from
their origin in t h e t a p e t a l c e l l s t o their d e p o s i t i o n o n
t h e p o l l e n greina and later ( 1972) d e s c r i b e d a l l u l t r a s t r u c t u r a ;
aapects o f pollen development f r o m t h e m i c r o s p o r e t e t r a d
s t a g e t o t h e three-celled s t a g e o f t h e p o l l e n g r a i n a n d
c o n f i r m e d t h e f i n d i n g s based o n t h e light m i c r o s c o p i c s t u d i e s
and a l s o c o r r e c t e d c e r t a i n e r r o n e o u s o b s e r v a t i o n s o f s o m e
earlier authors. D u n b a r (1973) s t u d y i n g t h e u l t r a s t r u c t u r e
and development o f pollen in t h e s a m e s p e c i e s c o n f i r m e d many
o f t h e o b s e r v a t i o n s made by
Carniel.Based o n t h e s e ultra-
s t r u c t u r a l f i n d i n g s , Strandhede(l973) g a v e a g e n e r a l i z e d
account o f t h e p o l l e n development o f
E. p a l u s t r i s .
Subsequently,
Meyer and Yaroshevskaya(1976) presented a b r i e f d e s c r i p t i o n
of t h e fine s t r u c t u r e o f the male g a m e t o p h y t e i n C a r e x
spicata, E. p a l u s t r i s , Eriophorum latifolium, R h y n c h o s p o r a
alba and
-
S c i r p u s sylvaticus. Based o n p o l l e n w a l l u l t r a s t r u c t u r e
they derived C y p e r a c e a e from the l i l i a c e o u a s t o c k t h r o u g h
Junacaceae. ~ l t h o u ~th e s t u d i e s in 1 9 7 0 s w e r e p r e d o m i n a n t l y
.,-
ultrastructure'oriented, a n o t a b l e light m i c r o s c o p i c c o n t r i -
bution w a s m a d e i n t h i s period by K r a l ( 1 9 7 1 ) w h o c o n d u c t e d
a c y t o t a x o n o m i c survey o f some N o r t h A m e r i c a n s p e c i e s o f
Abildqaardia, B u l b o s t y l 4 s and F i m b r i s t y l i s a n d r e p o r t e d
m i c r o s p o r o g e n e s i s i n s e v e r a l o f t h e s p e c i e s s t u d i e d but
failed t o o b s e r v e t h e c e l l plate
during cytokinesis in
m i c r o s p o r e m o t h e r cell.
Recently, Bir,Sidhu and Kamra(1906) s t u d i e d p o l l e n
m i t o s i s i n t e n . s p e c i e s , o n e each o f B u l b o s t y l i s a n d C a r e x ,
two o f S c i r p u s a n d s i x o f F i m b r i s t y l i s and u s e d t h e d a t a
for k a r y o t y p i c ana1ysis.Cayouette and M o r i s s e t ( 1 9 8 6 a , 1 9 8 6 b )
while m a k i n g a c y t o t a x o n o m i c s u r v e y o f s o m e N o r t h A m e r i c a n
S D ~ C ~0 f~ CBd r o v
r o n n ~ t r r lt h e
fnvmrt i n n n f
ttnrsnrlatnnrl
m i nnnsm..nnaa
in
C. asuatilis and C. recta. The previous embryological
work in Scleria and Carex with which the present work i s
concerned i s summarized in Table 1.
Table 1: Previous embryological work in Scleria
Species
Aspects studied
Scleria alata
Embryogeny and seed
and Carex
References
Schneider 1932
germination
S. biflora
-
Roxb.
Endothecium
Makde 1981a
S. foliosa
-
Rich.
Endosperm
Nijalingappa and Devaki
1979
Embryography
Meert
'&'
Goetghebeur 1979
Nijalingeppa & Arekal 1983
Embryology
Nijalingappa 1986
r& Td-
Embryogeny'
Shah & Neelakandan 1971
Endothecium
Makde 1981a
S. lithosperma
Endothecium
Makde 1981a
Hypostase
Makde 1981b
Obturator
Makde 1 9 8 2
Endosperm
Nijalingappa & Arekall983;
.
S. levis Retz,
hebecar a
K> Sw.
.
Makde &' Bhaskute 1 9 8 4
S . stockeiana
Boeck .
Endosperm
Nijalingappa & Arekal 1981
Endothecium .
Untawale 6 Bhasin 1973;
Makde 1 9 8 1 e
Species
Aspects studied
References
5 . tessellata
Pollen development
Tanaka 1941b
Endothecium
Makde 1981a
S. verrucosa
Willd.
Embryogeny
Juguet 1970e
undetermined
species
Embryo
Didrichsen 1897
Carex acuta L.
--
Pollen development
Juel 1900;
Willd.
Wulff 1939
C. aquatilis
-
Wg.
Embryo Sac
Jonsson
Pollen development
Stout 191 2
Microsporogenesis
Cayouette &
Morisset
C. arenaria
-
L.
Embryo Sac
1881
1986b
Hofmeister 1858
Guttenberg & Semlow 1957;
Juguet 1966b,1972;
.
Juguet b L e b e g u e l 9 6 6
C. aristata
C . atrata
-
R.Br.
L.
Megasporogenesis
Grezicka 1 9 6 4
Endosperm,
Embryogeny and
Seed germination
Schnei.der 1 9 3 2
C . capricornis
Meinshe
Megasprogenesis,
Embryo Sac
Tanaka 1 9 5 0
C . caryophyllacea
L a t.
P o l l e n development,
megasporogenesis,
embryo sac and
endosperm
Heilborn 1 9 1 8
Species
Aspects s t u d i e d
C, d i q i t a t a
-
L,
References
P o l l e n development
embryo sac and
endosperm
C. e r i c e t o r u m
-
H e i l b o r n 1918
P o l l e n development,
Poll.
megasporogenesis,
embryo sec and
C. f e d i a
--
Nees
(C.wallichiana
Presc)
endosperm
Heilborn
Embryology and seed
Shah 1959, 19628,
germination
1962b, 1 9 6 2 c , 1965
1967
'
C. f i l i c i n a
-
C. f l a v a
--
Nees
L,
,
1918
1968;
19818,
Endothecium
Makde
Microsporogenesis
B i r , S i d h u a n d Kamra 1986
Endothecium
Makde 1981a
Obturator
Makde 1982
P o l l e n development,
megasporogenesis,
embryo sac a n d
endoaperm-
Heilborn
C, q r a l l a t o r i a
Maxim,
P o l l e n development
l a n e k a 1939
C, q r a y i i
-
Embryo
D i d r i c h k e n 1894
Carey
1918
Endosperm, Embryo &
Seed g e r m i n a t i o n
S c h n e i d e r 1932
Species
Aspects s t u d i e d
C. h i r t a L .
--
C.
makinoensis
Franch.
References
Microsporogenesis
C a r n i e 1 1962
P o l l e n development
W i l l e 1886
Embryo Sac
H o f m e i s t e r 1858
Megasporogenesis &
Tanaka 1950
Embryo sac
C. m a r r o w i i
-
M e g a s p o r o g e n e s i s and
Boott
C. p a n i c e a
-
L.
Embryo sac
Tanaka 1950
Embryo Sac
Hofmeister
1858
P o l l e n development,
\
Megasporogenesis,
Embryo sac and
C. p e n d u l a
-
Huds.
endosperm
H e i l b o r n 191 8
Embryo
H o f m e i s t e r 1861
Polyembryony
M i r b e l 1810
( i n Braun 1860)
C, p i c t a
-
Steud.
P o l l e n development
Embryo sac
C. p i l u l i f e r a
-
C:
M a r t e n s 1939
P o l l e n development,
embryo sac a n d
C. p o d o q y n a
endosperm
H e i l b o r n 1918
Microsporogenesis
Tanaka 1941e
Franch.
C. p r a e c o x
-
S c h r e b . Embryo Sac
F i s c h e r 1800
Species
Aspects. s t u d i e d
C. r e c t a B o o t t
--
Microsporogenesis
References
C a y o u e t t e end
M o r i s s e t 19068.
C; s a l i n a
-
Wg.
Microsporogenesis
C a y o u e t t e and
M o r i s s e t t 1986a,
C. s p i c a t a
-
Huds.
P o l l e n development
Meyer and
Yaroshevskaya 1976
C, s u b s p a t h a c e a
Wormsk.
Microsporogeneais
C a y o u e t t e and
M o r i s s e t 1906a
C.
-
v e s i c a r i e L.
C. v u l p i n e
-
L.
Embryo s e c
Vesque 1 0 7 9
P o l l e n development
Wille 1 8 0 6
Embryo and s e e d
Undetermined
species
germination
D i d r i c h s e n 1897
P o l l e n development
W i l l e 1886