A Comparitive Study of Normalyoccuring Polypoloidy in Several Species of Allium
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r FORDHAM UNIVERSITY GRADUATE SCHOOL
..............................................
19,50.
This dissertation prepared under my direction by Jane Kins low M c E llig o tt
entitled
A
STUDY OP KfiRilALLY OCCURING
POLYPLOIDY IN SEVERAL SPECIES OP ALLIUM
has been accepted in partial fulfilment of the requirements for the
Degree o f
D
....................................
P r. C harles B erger (Faculty A dviser)
L
j
A COMPARATIVE STUDY OF NORMALLY OCCURRING POLYPLOIDY IN SEVERAL SPECIES OF ALLIUM
BY JANE K. MC ELLIGOTT B . A . , Marymount C o l l e g e *43 M . S . , Fordham U n i v e r s i t y *46
DISSERTATION SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN THE DEPARTMENT OF BIOLOGY AT FORDHAM UNIVERSITY New York 1 95 0
ProQuest Number: 10992989
All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is d e p e n d e n t upon the quality of the copy subm itted. In the unlikely e v e n t that the a u thor did not send a c o m p le te m anuscript and there are missing pages, these will be noted. Also, if m aterial had to be rem oved, a n o te will ind ica te the deletion.
uest ProQuest 10992989 Published by ProQuest LLC(2018). C opyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States C o d e M icroform Edition © ProQuest LLC. ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 4 8 1 0 6 - 1346
ill n
TABLE OF CONTENTS
LIST OF FIGURES.............................................
...
ACKNOWLEDGEMENT..........................
Page lv xi
I .
INTRODUCTION.......................................................
1
II.
MATERIAL AND METHODS . . . ............................
6
III.
OBSERVATIONS AND RESULTS..........................
10
A llium A llium A llium A llium A llium A llium A llium A llium
s t e l l a t u m . , .................................. 11 fIstulosum • • 13 s c h o e n o p r a s u m ................... 2? a n g u l o s u m ....................................... 43 v a l l d u m ............................................ 67 p o r r u m .............................................. 68 ammophllum. . . . . ................ 86 tuberosum . ............................102
IV.
DISCUSSION....................
129
V.
SUMMARY.....................................
138
BIBLIOGRAPHY............................................................................ 141
L
.J
lv r
n LIST OF FIGURES
Figure 1. 2.
3.
4.
5.
6. 7,
8.
9. 10, 11.
12,
13.
Page
Drawing o f an A l l i u m s e e d l i n g s h o w in g t h e v a r i o u s t i s s u e s , . . . ..............
9
P h o to m icro g rap h of a d i p l o i d c e l l o f Allium s t e l l a t u m sh o w in g f o u r t e e n p r o m e t a p h a s e c h r o m o s o m e s , ....................................
15
P ho to m icrog rap h of a p a i r e d p o l y p lo i d p r o p h a s e o f A l l i u m s t e l l a t u m sh ow in g r e l a tio n al c o ilin g , ............... .............................. . ............
15
Photom icrograph of a l a t e p o ly p lo id prophase o f A l l i u m s t e l l a t u m sh o w in g p a i r i n g and r e l a t i o n a l c o i l i n g ................................................. • • • • •
1?
P h o to m ic ro g ra p h of a d i p l o i d m etaphase of A l l i u m s t e l l a t u m sho w in g v e r y c o n t r a c t e d c h r o m o s o m e s . . . ................. ......................................, , • • • •
17
P h oto m icrog raph of a p o l y p lo i d anaphase of A l l i u m s t e l l a t u m .......... ...............................
19
P h o t o m i c r o g r a p h o f a l o n g i t u d i n a l s e c t i o n of a young s e e d l i n g 13 mm. l o n g o f A l l i u m s t e l l a t u m sh ow in g t h e v a r i o u s t i s s u e s . ..........
21
Photom icrograph of a b i n u c l e a t e c e l l of A l l i u m s t e l l a t u m s h o w in g a l a r g e n u c l e u s and a m icronucleus. .................................................
23
P hotom icrograph of a c e l l of A llium s t e l l a tum w i t h two m l c r o n u c l e l ..............................................
23
P h o to m icro g rap h of a d u m b -b ell shaped n u c l e u s o f A l l i u m s t e l l a t u m ..........................
25
P h o to m icro g rap h of a c e l l of A llium s t e l l a tum w i t h t e l o p h a s e b r i d g e s and chro m osomal f r a g m e n t s ................
25
P h o to m ic ro g ra p h of a d i p l o i d c e l l of A llium f i s t u l o s u m sho w in g t h e k a r y o t y p e and s a t e l l i t e s .....................................................................
29
P h o t o m i c r o g r a p h o f d i p l o i d and t e t r a p l o i d r e s t i n g n u c l e i of A llium f I s t u l o s u m
29
V
n
Figure 14.
P h o t o m i c r o g r a p h o f a p o l y p l o i d p r o p h a s e of A l l i u m f l s t u l o s u m s h ow ing p a i r i n g and r e l a t i o n a l c o i l i n g * ..........................................• • • • • • . .
31
15.
Photom icrograph of a very c o n tr a c te d m eta p h a s e o f A l l i u m f l s t u l o s u m . . . . .......................................... 31
16,
P h o t o m i c r o g r a p h o f a p o l y p l o i d m e t a p h a s e of A l l i u m f l s t u l o s u m sh o w in g t h i r t y - t w o ..................... chromosomes
33
P h o to m ic ro g ra p h of a p o l y p l o i d anaphase of A l l i u m f l s t u l o s u m * ................................................
33
P h o t o m i c r o g r a p h o f a p o l y p l o i d t e l o p h a s e of A l l i u m f l s t u l o s u m .............................
35
P h o t o m i c r o g r a p h of a l o n g i t u d i n a l s e c t i o n o f a young s e e d l i n g 20 mm. l o n g of A l l i u m f l s t u l osum s h o w in g t h e v a r i o u s t i s s u e s . .........................
37
Ph oto m icro grap h of a d u m b -bell shaped n u c le u s .................................. of Allium f l s t u l o s u m
39
P h o t o m i c r o g r a p h of a b i n u c l e a t e c e l l o f A l l i u m f l s t u l o s u m s h o w in g t h e l a r g e n u c l e u s and t h e m i c r o n u c l e u s .........................................................................
39
P ho tom icro grap h of a d i p l o i d t e lo p h a s e of A l l i u m f l s t u l o s u m sh ow ing chro m osomal b r i d g e s a n d c hrom oso m a l f r a g m e n t s ...................
41
P h o t o m i c r o g r a p h of a d i p l o i d c e l l of A l l i u m s c h o e n o p r a s u m s h o w in g t h e k a r y o t y p e . ....................
45
P h o to m icro g rap h of a c e l l of A llium schoeno p r a s u m s h o w in g a s a t e l l i t e ................
45
P h o t o m i c r o g r a p h o f a d i p l o i d and a t e t r a p l o l d c e l l o f A l l i u m s c h o e n o p r a s u m . . .....................
47
17. 18. 19.
20. 21.
22.
23. 24. 25. 26.
27.
L
Page
P h o t o m i c r o g r a p h o f a p o l y p l o i d p r o p h a s e of A l l i u m s c h o e n o p r a s u m sh o w in g p a i r i n g a n d rela tio n al c o llin g .......................... Photom icrograph of a p o ly p lo id m id-prophase o f A l l i u m s c h o e n o p r a s u m show ing p a i r i n g and r e l a t i o n a l c o i l i n g ...................
« . . . 47
49
vi r
-i Figure 28.
29.
Page P h o t o m i c r o g r a p h o f an e a r l y a n a p h a s e o f A l l i u m s c h o e n o p r a s u m show in g t h i r t y - t w o .............................................. chromosomes
49
P h o to m ic ro g ra p h of a p o l y p l o i d t e l o p h a s e of A l l i u m s c h o e n o p r a s u m show in g n um erou s c hro m o som a l b r i d g e s ... ................................................
51
30.
P h o t o m i c r o g r a p h o f a l o n g i t u d i n a l s e c t i o n of a young s e e d l i n g o f A l l i u m s c h o e n o p r a s u m 18 mm. l o n g sh o w in g t h e v a r i o u s t i s s u e s ..................... 53
31.
P h o to m icro g rap h of a d u m b -b ell shaped n u c le u s i n a c e l l o f A l l i u m s c h o e n o p r a s u m ............................... 55
32.
Ph otom icrog raph of a d i p l o i d te lo p h a s e of A l l i u m s c h o e n o p r a s u m show ing c h rom oso m a l b r i d g e s an d f r a g m e n t s . . . .............................................
55
P h o to m ic ro g ra p h of an e a r l y anaphase of A l l i u m a n g u lo s u m s ho w in g a s m a l l s a t e l l i t e . . . .
58
33. 34.
P h o t o m i c r o g r a p h o f a d i p l o i d and a t e t r a p l o i d c e l l o f A l l i u m a n g u l o s u m ................................................... 58
35.
P h o t o m i c r o g r a p h o f a p o l y p l o i d p r o p h a s e of A l l i u m a n g u lo s u m sh o w in g p a i r i n g and r e l a t i o n a l c o i l i n g . . .................................
60
P hotom icrograph of a p o ly p lo id m etaphase of A l l i u m a n g u l o s u m ........................
60
P h o to m icro g rap h of a p o ly p lo id e a r l y anaphase of Allium angulosum . ............
62
36. 3?. 38.
Photom icrograph of a p o ly p lo id l a t e anaphase of Allium angulosum . ........................................................ 62
39.
P h o t o m i c r o g r a p h of a b l n u c l e a t e c e l l o f A l l i u m a n g u lo s u m sho w in g a l a r g e n u c l e u s and a m i c r o n u c l e u s ........................................................................... 64
40.
P h o t o m i c r o g r a p h o f a d i p l o i d t e l o p h a s e of A l l i u m a n g u lo s u m show in g chro m osomal b r i d g e s and f r a g m e n t s .....................................................
64
P h o to m ic ro g ra p h of a d u m b -b e ll shaped n u c le u s o f A l l i u m a n g u l o s u m ...........................................
66
41.
v il r
“l
Figure 42.
L
Page
P h o to m ic ro g ra p h of a d i p l o i d c e l l of Allium v a l i d u m shov/ing a s u b t e r m i n a l s p i n d l e a t t a c h m e n t r e g i o n .................
70
45.
P h o t o m i c r o g r a p h o f a d i p l o i d and t e t r a p l o i d r e s t i n g c e l l s o f A l l i u m v a l i d u m .............................., 7 0
44.
P h o t o m i c r o g r a p h o f a p a i r e d and r e l a t l o n s l l y c o i l e d e a r l y p r o p h e s e of A l l i u m v a l i d u m .....................72
45.
P h o t o m i c r o g r a p h o f a p a i r e d and r e l a t i o n a l l y c o i l e d m i d - p r o p h a s e o f A l l i u m v a l i d u m ...................... 72
46*
Pho to m icro grap h of a p o ly p lo id prom etaphase of* A l l i u m v a l i d u m sho w in g r e l a t i o n a l c o l l i n g a n d p a i r i n g . . .......................................................
74
47.
P h o to m ic ro g ra p h o f a p o l y p l o i d m etaphase of A l l i u m v a l i d u m show in g a wide s e p a r a t i o n of t h e c h r o m a t i d ' s ............ .......................................... . . . .............. 74
48.
P h o t o m i c r o g r a p h o f a p o l y p l o i d a n a p h a s e of A l l i u m v a l i d u m .............................
76
49.
Photom icrog rap h of a p o ly p lo id m etaphase of A l l i u m v a l i d u m sh ow in g t h e c h r o m a t i n l u m p ................76
50.
Photom icrograph of a l o n g i t u d i n a l s e c tio n o f a young s e e d l i n g o f A l l i u m v a l i d u m 15 mm. l o n g sh o w in g t h e v a r i o u s t i s s u e s .................................. 78
51.
P ho tom icrog raph of a b i n u c l e a t e c e l l of A l l i u m v a l i d u m show in g t h e l a r g e n u c l e u s and t h e m i c r o n u c l e u s ........................
80
52.
Photom icrograph of a v ery l a t e t e lo p h a s e of A l l i u m v a l i d u m sh o w in g t h e f o r m a t i o n o f a m i c r o n u c l e u s ................................................................................... 80
55.
P h o t o m i c r o g r a p h o f a t e l o p h a s e of A l l i u m v a l i d u m s h o w in g b r i d g e s and f r a g m e n t s ...................... 82
54.
Photom icrograph of a dum b-bell shaped n u cleu s o f A l l i u m v a l i d u m . . . , ............................................................. 84
55.
P h o to m ic ro g ra p h of a d i p l o i d c e l l o f A llium po r r u m sh o w in g t h e c o n f i g u r a t i o n o f t h e c h r o m o s o m e s .........................................
88 -J
v ili
Figure 56.
57. 58.
59.
60.
61. 62.
63.
64.
65. 66.
Page P h o t o m i c r o g r a p h o f a d i p l o i d c e l l of A l l i u m p o rr u m sh o w in g a l a r g e s a t e l l i t e a t t a c h e d t o t h e s h o r t e r arm of a chro m osom e..................... ..
88
P h o t o m i c r o g r a p h o f a d i p l o i d c e l l and a t e t r a p l o i d c e l l o f A l l i u m p o r r u m ...............................
90
P hotom icrograph of a p o ly p lo id prophase of A l l i u m p o r r u m s h o w in g p a i r i n g and r e l a t i o n a l c o l l i n g . . . .................................
90
P h o to m ic ro g ra p h o f a m id -p ro p h a s e of A llium p o r r u m show in g p a i r i n g and r e l a t i o n a l c o l l i n g .........................................
92
P h o to m ic ro g ra p h of a p o l y p l o i d m etaphase o f A l l i u m p o r r u m sh o w in g s i x t y - f o u r c h ro m o somes w i t h w i d e l y s p r e a d c h r o m a t i d s ......................
92
P h o t o m i c r o g r a p h o f a p o l y p l o i d a n a p h a s e of A l l i u m p o r r u m ............................
94
P h o t o m i c r o g r a p h o f a l o n g i t u d i n a l s e c t i o n of a y o u n g s e e d l i n g 25 mm. l o n g o f A l l i u m porrum shoxtflng t h e v a r i o u s t i s s u e s ..........................................
96
Photom icrograph of a b in u c l e a t e c e l l of A l l i u m p o r r u m show in g a l a r g e n u c l e u s and a m i c r o n u c l e u s ..............................................................................
98
P h o to m icro g rap h of a t e l o p h a s e of A llium p o r r u m s h o w in g c h rom o so m a l b r i d g e and f r a g m e n t s . . . . .....................................................................
98
P h o to m icro g rap h of a dum b-bell shaped n u cleu s o f A l l i u m p o r r u m .....................................................................
100
P h o t o m i c r o g r a p h o f a d i p l o i d n u c l e u s and a t e t r a p l o i d n u c l e u s o f A l l i u m ammophllum. . . .
104
67.
Photom icrograph of a p o ly p lo id e a r ly prophase A l l i u m ammophllum show ing p a i r i n g and r e l a t i o n a l c o l l i n g ................................................................. 104
68.
Photom icrograph of a p o ly p lo id m id-prophase A l l i u m ammophllum s h o w in g r e l a t i o n a l c o i l i n g and p a i r i n g .............................................
106
ix
Figure 69.
70.
Page P h o t o m i c r o g r a p h o f a p o l y p l o i d m e t a p h a s e of A l l i u m ammophilum sh o w in g s i x t y - f o u r c h ro m o somes a n d k a r y o t y p e ...................................
106
P h o t o m i c r o g r a p h o f a p o l y p l o i d m e t a p h a s e of A l l i u m ammophllum sho w in g s i x t y - f o u r g r e a t l y c o n t r a c t e d c h r o m o s o m e s . . . . ..............................................
108
71.
P h o to m icro g rap h o f a p o l y p l o i d anaphase of A l l i u m ammophllum.................................................................... 108
72.
P h o t o m i c r o g r a p h o f a l o n g i t u d i n a l s e c t i o n of a yo ung s e e d l i n g 18 mm. l o n g o f A l l i u m ammophllum sh ow in g t h e v a r i o u s p a r t s ...................
110
P h otom icrograph o f a d i p l o i d te lo p h a s e of A l l i u m ammophllum sh o w in g chro m osom al b r i d g e s an d f r a g m e n t s ..................................................
112
73.
74.
P h otom icrograph of a b i n u c l e a t e c e l l of A l l i u m ammophllum s h o w in g a l a r g e n u c l e u s and a m i c r o n u c l e u s . . . . ........................................................ 112
75.
P h o t o m i c r o g r a p h o f a d i p l o i d m e t a p h a s e of A l l i u m t u b e r o s u m s h o w in g t h e c o n f i g u r a t i o n and w i d e l y s p r e a d c h r o m a t i d s .................... 116
76.
P h o t o m i c r o g r a p h o f a d i p l o i d c e l l and a t e t r a p l o i d c e l l o f A l l i u m t u b e r o s u m . . . . ..............
116
77.
P h o t o m i c r o g r a p h o f an e a r l y p o l y p l o i d p r o p h a s e o f A l l i u m t u b e r o s u m sho w in g p a i r i n g and r e l a t i o n a l c o i l i n g ........................................................ 118
78.
Photom icrograph of a l a t e p o ly p lo id prophase o f A l l i u m t u b e r o s u m show in g r e l a t i o n a l c o i l i n g and p a i r i n g ...............................................................
118
79.
P h o t o m i c r o g r a p h o f a p o l y p l o i d m e t a p h a s e of A l l i u m t u b e r o s u m showring t e t r a c h r o m o s o m e s . . . . 120
80.
P h o t o m i c r o g r a p h o f a p o l y p l o i d m e t a p h a s e of A l l i u m t u b e r o s u m i n w h i c h some o f t h e s p i n d l e a tta c h m e n t r e g i o n s have s e p a r a t e d
81.
120
Photom icrograph of a p o ly p lo id anaphase of A l l i u m t u b e r o s u m ....................................................................... 122
~ i
X
* ,
F i g . 14
F ig . 15
_i
32
E x p la n a tio n of F ig u re s 16,
P h o t o m i c r o g r a p h o f a p o l y p l o i d m e t a p h a s e of A l l i u m f i s t u l o s u m t a k e n from an a c e t o - o r c e i n sm ear p r e p a r a t i o n sh o w in g t h i r t y - t w o c h ro m o so m es.
M a g n if ic a tio n x 1400.
«
17.
Photom icrograph of a p o ly p lo id anaphase of A l l i u m f l s t u l o s u m t a k e n fro m a n a c e t o - o r c e i n sm e ar p r e p a r a t i o n .
l.
M a g n i f i c a t i o n x 1400.
33
*
F i g . 16
F ig . 17
E x p la n a tio n of F ig u re Photom icrograph of a p o ly p lo id te lo p h a s e o f A l l i u m f i s t u l o s u m t a k e n f r o m an a c e t o o r c e in smear p r e p a r a t i o n . x 1400.
M agnification
35
n
r
F i g . 18
36 r
i
E x p la n a tio n of F igure 19. Photom icrograph of a l o n g i t u d i n a l s e c t i o n o f a y o un g s e e d l i n g t w e n t y m i l l i m e t e r s l o n g o f A l l i u m f i s t u l o s u m sh o w in g t h e v a r i o u s tissu es.
L
M a g n i f i c a t i o n x 175. EGT =
ectoderm is
FL =
first
RT =
re g io n of t r a n s i t i o n
C
co tyledon
=
leaf
CC =
c o rte x of cotyledon
CT =
c o rte x of re g io n of t r a n s i t i o n
37 n
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F i g . 19
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E xplanation of F igures 20. P h o to m ic ro g ra p h of a d u m b -b ell shaped n u c le u s A l l i u m f l s t u l o s u m t a k e n fro m an a c e t o o r c e l n sm e ar p r e p a r a t i o n .
M agnification
x 14Q0j 2 1. P h o t o m i c r o g r a p h o f a b i n u c l e a t e c e l l o f A l l i u m f i s t u l o s u m r sh o w in g t h e l a r g e n u c l e u s . ■ "
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An a c e t o - o r c e l n sm ear p r e p a r a t i o n . M a g n i f i c a t i o n x 14Q0#
L .
39 n
Fig.
L
21
40
r
n
E x p la n atio n of Figure 22. P h o t o m i c r o g r a p h o f a d i p l o i d t e l o p h a s e o f A l l i u m f i s t u l o s u m s ho w ing c h ro m o s o m a l b r i d g e s and chro m o so m al f r a g m e n t s . sm e a r p r e p a r a t i o n .
L
An a c e t o - o r c e i n
M a g n if ic a tio n x 1400.
41 ~ i
F i g . 22
L
42 r~
-i
and two w i t h s u b - t e r m i n a l s p i n d l e a t t a c h m e n t r e g i o n s (fig .
23).
The l a t t e r h a v e a s m a l l s a t e l l i t e w h i c h i s
a t t a c h e d t o t h e s h o r t e r arm by a l o n g f i l a m e n t ( f i g .
24).
T h i s i s i n a g r e e m e n t w i t h t h e work o f Levan ( 1 9 3 2 ) .
The
chromosomes a r e s m a l l ,
s h o r t and s l e n d e r .
The y oung s e e d l i n g s o f t h i s s p e c i e s i n f r e c i u e n t l y f o rm ed t e t r a p l o i d c e l l s . When p o l y p l o i d y d i d o c c u r ,
T hese u n d e r w e n t n o r m a l m i t o s i s . i t d i d so i n c e l l s w h ic h
were much l a r g e r t h a n t h e d i p l o i d c e l l s
(fig .
25).
The
chromosomes w ere s e e n t o be c l o s e l y p a i r e d and r e l a t i o n a l l y c o i l e d t h r o u g h o u t p r o p h a s e ( f i g s . . 26 and 2 7 ) . A l t h o u g h t h e s p i n d l e a t t a c h m e n t r e g i o n s o f t h e c h ro m o somes were n o t a c t u a l l y s e e n t o r e m a i n s i n g l e and undivided u n t i l m etaphase,
it
i s p o s s i b l e t h a t such a
s i t u a t i o n o b t a i n s s in c e t h i s i s th e e s s e in the o t h e r sp e c ie s .p rev io u sly d e sc rib e d . fo llo w ed the normsl p a t t e r n .
Anaphas e s e p a r a t i o n O c c a s i o n a l l y , however,
p o l y p l o i d t e l o p h a s e s were s e e n w i t h num erous c h ro m o so m a l b r i d g e s ( f i g .
29).
P o l y p l o i d d i v i s i o n s were f ou n d t o be p r e s e n t i n t h e c o r t e x o f t h e c o t y l e d o n and o f t h e t r a n s i t i o n r e g i o n b e t w e e n r o o t and s h o o t .
A few p o l y p l o i d c e l l s
we re s e e n i n t h e e c t o d e r m i s o f t h e c o t y l e d o n ( f i g . 3 0 ) . The p o l y p l o i d c e l l s we re f o u n d i n t h e b a s a l p o r t i o n of the c o ty le d o n . P o l y p l o i d d i v i s i o n s we re f i r s t L
observed in _J
s e e d l i n g s 10 mm. l o n g . evident.
At 33 mm. no p o l y p l o i d y was
D i v i s i o n s o f any k i n d were l a c k i n g e x c e p t i n
the m erlstem of the f i r s t w ere p r e s e n t .
l e a f where d i p l o i d d i v i s i o n s
The f i r s t l e a f a p p e a r e d a t a p p r o x i m a t e l y
40 mm. I n t h e same r e g i o n w i t h t h e p o l y p l o i d c e l l s , blnucleate c e l ls ,
dum b-bell shaped n u c l e i ( f i g . 3 2 ),
t e l o p h a s e b r i d g e s and ch ro m o s o m a l f r a g m e n t s ( f i g . 32} w ere f o u n d . None o f t h e s e a b e r r a t i o n s a p p e a r e d w i t h g r e a t f r e q u e n c y a l t h o u g h t h e chro m o som a l b r i d g e s and f r a g m e n t were more common t h a n t h e o t h e r tw o. A l l i u m e n g u lo s u m L . A l l i u m a n g u l o s u m . commonly known a s Mouse G a r l i c , i s n a t i v e t o S i b e r i a and c o n s i s t s o f one b u l b o r two bulbs to g e th e r a tta c h e d to a h o r i z o n ta l or ascending rhizom e.
Many v i o l e t f l o w e r s a r e b o r n e on an u m b e l .
T h i s s p e c i e s c o n s t i t u t e s a t y p i c a l e xam ple o f p o ly p lo id y w ith in the s p e c i e s .
Levan ( 1 9 3 5 ) h a s among
h i s c u l t u r e s f o r m s w i t h a d i p l o i d number o f 16 a s w e l l a s t h o s e w i t h a d i p l o i d number o f 1 0 0 .
The p a r t i c u l a r
one u s e d i n t h i s i n v e s t i g a t i o n h a s a d i p l o i d number o f s i x t e e n and a l l chromos omes a r e m e d i a n i y i n s e r t e d (Levan, 1 9 3 5 ). sa tellite
One p a i r o f chromosomes b e a r s a s m a l l
( f ig . 33).
44 r
E x p la n a tio n of F ig u r e s 23. P h o to m ic ro g ra p h of a d i p l o i d c e l l of A llium s c h o e n o p r a s u m t a k e n fro m an a c e t o - n i g r o s l n e sm ear p r e p a r a t i o n sh o w in g t h e k a r y o t y p e . M a g n i f i c a t i o n x 1350* 24. P h o to m ic ro g ra p h of a d i p l o i d c e l l of A llium s c h o e n o p r a s u m t a k e n from a n a c e t o - n i g r o s i n e sm ear p r e p a r a t i o n s h o w in g a s a t e l l i t e . M a g n i f i c a t i o n x 14Q0*
L.
45
F l S.
F ig . 24 L
46 r
i
E x p lan atio n of F ig u re s 2 5 . P h o t o m i c r o g r a p h o f a d i p l o i d and a t e t r a p l o i d c e l l of Allium schoenoprasum ta k e n fr o m an a c e t o - n i g r o s i n e sm ear p r e p a r a t i o n . M a g n ific a tio n x 1400. 26. P h o to m icro g rap h of a p o l y p l o i d p ro p h a s e of t
A l l i u m s c h o e n o p r a s u m t a k e n fro m an a c e t o o r c e l n sm e ar p r e p a r a t i o n s h ow in g p a i r i n g and r e l a t i o n a l c o l l i n g . 1400.
M ag n lficatio n x
4?
F ig . 26
48 r
E x p la n a tio n of F ig u re s 2?. P h otom icrograph of a p o l y p lo i d p ro p h a se of A l l i u m s c h o e n o p r a s u m show in g p a i r i n g and re la tio n a l co ilin g . preparation.
An a c e t o - o r c e i n sm ear
M a g n if ic a tio n x 1400.
2 8 . P h o t o m i c r o g r a p h o f an e a r l y a n a p h a s e o f A l l i u m s c h o e n o p r a s u m t a k e n fro m an a c e t o nigrosine
sm ear p r e p a r a t i o n s h o w in g t h i r t y
two c h ro m o s o m e s.
L
M a g n i f i c a t i o n x 1250*
49 “i
F i g . 2?
E x p la n atio n of Figure . Photom icrograph of a p o l y p lo i d te l o p h a s e of A llium schoenoprasum ta k e n from an a c e t o n l g r o s i n e sm ear p r e p a r a t i o n s h o w in g numerou c hro m o som a l b r i d g e s .
M a g n ific a tio n x 1400.
E x p la n atio n of Figure P hotom icrograph of a l o n g i t u d i n a l s e c t i o n o f a y o un g s e e d l i n g 18 mm. l o n g o f A l l i u m s c h o e n o p r a s u m s ho w ing t h e v a r i o u s t i s s u e s . M a g n i f ic a t i o n x 175. ECT = e c t o d e r m i s FL =
f i r s t leaf
HT =
reg ion of t r a n s i t i o n
C
cotyledon
=
CC =
c o r t e x of c o ty le d o n
CT =
c o r te x of r e g io n of t r a n s i t i o n
•'
v •
• ; * *
4
‘
'** j. T , ^ *7
■
-
'
F i g . 30
^
54 r
E x p la n atio n of F ig u re s 31.
P h o to m icro g rap h of a dum b-bell shaped n u c le u s t a k e n f r o m an a c e t o - o r c e i n s m e a r p r e p a r a t i o n of A llium schoenop rasum .
32.
M a g n i f i c a t i o n x 14Q0.
P h o to m ic ro g ra p h of a d i p l o i d t e l o p h a s e of A lliu m s c h o e n o p r a s u m t a k e n fro m an a c e t o - n l g r o s i n e sm ear p r e p a r a t i o n s h o w in g c h ro m o s o m a l b r i d g e s and f r a g m e n t s .
L.
M a g n if ic a tio n x 1400.
55 n r
F i g . 31
F ig . 32 _i L
56 n
An i n v e s t i g a t i o n o f t h e t r a n s i t i o n r e g i o n and c o t y l e d o n o f t h e y o ung s e e d l i n g r e v e a l e d t e t r a p l o i d y t o be a b u n d a n t . d ip lo id
(fig .
These c e l l s were l a r g e r t h a n t h e
5 4 ) and u n d e r w e n t n o r m a l m i t o s i s
( f i g s . 3 5 , 3 6 , 3 7 , and 3 8 ) .
Only p a i r e d p o l y p l o i d
p r o p h e s e s were f o u n d i n d i c a t i n g t h a t t h e p o l y p l o i d c e l l s undergo j u s t a s i n g l e d i v i s i o n . osomes we re r e l a t i o n a l l y
co iled .
P o l y p l o i d d i v i s i o n s w ere f i r s t s e e d l i n g s 10 mm. l o n g .
The p a i r e d ch ro m
observed in
Since s u f f i c i e n t m a t e r i a l
c o u l d n o t be o b t a i n e d t o c o m p l e t e t h e i n v e s t i g a t i o n , i t c a n n o t be d e f i n i t e l y
s a i d a t w hat l e n g t h p o l y
p l o i d y d i s a p p e a r e d n o r i n what r e g i o n s i t was f o u n d . S i n c e A l l i u m a n g u lo s u m seems t o f o l l o w t h e p a t t e r n s e t by o t h e r members of t h i s g e n u s h a v i n g a d i p l o i d chromosome number o f s i x t e e n ,
i t m i g h t be assum ed t h a t
t h e t e t r a p l o i d c e l l s c e a s e d t o be e v i d e n t a t a p p r o x i m a t e l y 40 mm.
A l s o t h a t t h e y were p r e s e n t i n t h e
c o r t e x o f t h e c o t y l e d o n and o f t h e r e g i o n of t r a n s i t i o n and p o s s i b l y i n t h e e c t o d e r m i s . A8 i n A l l i u m f i s t u l o s u m and A l l i u m s c h o e n o p r a s u m . b lnucleate c e lls
( f i g . 3 9 ) w ere f o u n d i n t h e same r e g i o n
a s t h a t i n w h i c h t h e t e t r a p l o i d c e l l s w e re f o u n d . Chromosomal b r i d g e s ,
f r a g m e n t s ( f i g . 4 0 ) and d u m b - b e l l
s h a p e d n u c l e i ( f i g . 4 1 ) w ere a l s o o b s e r v e d .
L
5? r
i
E x plsn atio n of F igures 55.
P h o t o m i c r o g r a p h o f an e a r l y a n a p h a s e o f A l l i u m a n g u lo s u m t a k e n f r o m an a c e t o - o r c e i n sm ear p r e p a r a t i o n sh ow in g a s m a l l s a t e l l i t e . M a g n ific a tio n x 1400.
54.
P h o t o m i c r o g r a p h o f a d i p l o i d and a t e t r a p l o i d c e l l o f A l l i u m a n g u lo s u m t a k e n f r o m an a c e t o o r c e i n sm e ar p r e p a r a t i o n .
L
M a g n if ic a tio n x 1400.
58
59 r
i
E xplanation of Figures 35.
P h o t o m i c r o g r a p h o f a p a i r e d and r e l a t i o n a l l y c o i l e d p o l y p l o i d p r o p h a s e o f A l l i u m a n g u lo s u m t a k e n fr om a n a c e t o - o r c e i n sm e ar p r e p a r a t i o n . M a g n if ic a tio n x 1400.
36.
Photom icrograph of a p o ly p lo id m etaphase of A l l i u m a n g u l o s u m t a k e n from an a c e t o o r c e i n sm ear p r e p a r a t i o n .
M ag n ificatio n
x 1400.
L
_i
60
F i g . 35
Fig.
36
Explanation o f Figures 37.
Photom icrograph of a p o ly p lo id e a r l y ana p h a s e o f A l l i u m a n g u lo s u m t a k e n fro m an a c e t o - o r c e i n sm ear p r e p a r a t i o n . M a g n i f i c a t i o n x 1400*
38.
P h o to m icro g rap h of a p o l y p l o i d l a t e a n a p h a s e o f A l l i u m a n g u lo s u m t a k e n fro m an a c e t c - o r c e l n sm ear p r e p a r a t i o n . M a g n i f ic a t i o n x 1400,
L
62
F ig . 38 _i L
63 r
E xplanation of F igures 39.
P h otom icrog raph of a b i n u c l e a t e c e l l of A llium a n g u lo s u m t a k e n fr om an a c e t o - o r c e l n sm ear p r e p a r a t i o n show ing a l a r g e n u c l e u s and a m icronucleus.
40.
M a g n ific atio n x 1400.
P h o t o m i c r o g r a p h o f a d i p l o i d t e l o p h a s e of A l l i u m a n g u lo s u m sh o w in g c h ro m o s o m a l b r i d g e s and f r a g m e n t s . paratio n .
L
An a c e t o - o r c e i n sm e ar p r e
M ag n ificatio n x 1400.
_ i
64
F ig . 40
L
E x p la n atio n of Figure P ho to m icro g rap h of a dum b-bell shaped n u c l e u s o f A l l i u m a n g u lo s u m t a k e n fro m an a c e t o - o r c e i n sm e ar p r e p a r a t i o n . M a g n ific a tio n x 1400.
F i g . 41
A l l i u m v&lldum W a t s . A l l i u m v & ll d u m . known a s t h e P a c i f i c On io n , belongs to the ornam ental o n io n s.
It
i s n a tiv e to
N o r t h America and grows i n l a r g e c lu m p s one t o two f e e t high w ith la rg e c l u s t e r s
of l a v e n d e r f l o w e r s .
The d i p l o i d number r e p o r t e d by Levan ( 1 9 3 2 ) is tw en ty -eig h t.
The chromosom es a r e l a r g e .
Twenty-
f o u r h a ve m e d i a n s p i n d l e a t t a c h m e n t r e g i o n s and f o u r have s u b - t e r m i n a l s p i n d l e a tta c h m e n t r e g i o n s ( f i g . 4 2 ) . Two p a i r s o f t h e c hro m o s o m e s, t h o s e w i t h t h e s h o r t a r m s , have v ery s m a ll s a t e l l i t e s
a t t a c h e d t o them ( L e v a n ,
1932). As i n t h e p r e v i o u s l y d e s c r i b e d s p e c i e s ,
i t was
f o u n d t h a t t h e yo u ng s e e d l i n g s r e g u l a r l y and v e r y f r e q u e n t l y f orm ed t e t r a p l o i d c e l l s . d i v is io n s occurred in la rg e c e l l s
Such p o l y p l o i d
( f ig . 43).
The
chromosom es w ere s e e n t o be c l o s e l y p a i r e d and r e l a t l o n a l l y c o i l e d from e a r l y p ro p h a s e to m etaphase (fig s.
44, 4 5 , 4 6 , and 4 7 ) .
At m e t a p h a s e ( f i g .
t h e p a i r e d chromosomes s e p a r a t e d .
47),
T h i s was f o l l o w e d
by a n o t h e r s e p a r a t i o n w h i c h marked t h e b e g i n n i n g o f a n a p h a s e m ovem ent. were n o r m a l .
An a p h ase ( f i g . 4 8 ) and t e l o p h a s e
No o c t o p l o i d c e l l s were s e e n .
A c c o r d i n g t o Levan ( 1 9 3 2 ) , a f r a g m e n t e d c h ro m o some i s f o u n d i n A l l i u m v a l i d u m .
One o f t h e l o n g
armed chromosomes b r o k e n e a r t h e c o n s t r i c t i o n .
One p a r t
o f t h i s chromosome d e v e l o p e d i n t o a n o r m a l chromosome and t o o k i t s p l a c e on t h e m e t a p h a s e p l a t e w h i l e t h e o t h e r p a r t of i t was l i k e a lump o f c h r o m a t i n .
T his
may be s e e n i n f i g u r e 4 9 . P o l y p l o i d d i v i s i o n s were f o u n d t o be p r e s e n t i n t h e c o r t e x o f t h e c o t y l e d o n and o f t h e t r a n s i t i o n r e g i o n b e t w e e n r o o t and s h o o t ( f i g . were se en i n th e e c t o d e r m i s .
50).
A few p o l y p l o i d c e l l s
The b a s a l p o r t i o n o f t h e
c o t y l e d o n c o n t a i n e d many more p o l y p l o i d c e l l s
than
t h a t re g io n c l o s e r to the seed . P o l y p l o i d d i v i s i o n s were f i r s t
seen in s e e d l i n g s
10 mm. l o n g and t h e y p e r s i s t e d u n t i l t h e s e e d l i n g m e a s u r e d 45 mm.
Here o n l y a few d i v i s i o n s were s e e n
and t h o s e w e re p r e s e n t i n t h e m e r i s t e m a t i c r e g i o n .
The
f i r s t l e a f emerged from t h e c o t y l e d o n when t h e s e e d l i n g m e a s u r e d from 60 t o 70 mm. C e r t a i n a b n o r m a l i t i e s were a l s o f o u n d t o e x i s t i n t h e same r e g i o n a s t h e p o l y p l o i d c e l l s . cells
(fig .
(fig .
5 5 ) , and d u m b - b e l l s h a p e d n u c l e i ( f i g .
e v id en t.
51),
B lnucleate
t e l o p h a s e b r i d g e s and f r a g m e n t s 5 4 ) were
A l l t h r e e of t h e s e were f r e q u e n t l y o b s e r v e d . A l l i u m p o rru m L.
A l l i u m p o r r u m i s known t o t h e l a y p e r s o n a s t h e common l e e k .
Although i t
i s n a t i v e t o t h e s h o r e s of
69 r
E x planation of Fig u res 42.
P h o to m icro g rap h of a d i p l o i d c e l l of Allium v a l l d u m t a k e n f r o m an a c e t o - n l g r o s l n e
smesr
p r e p a r a t i o n sh ow in g a s u b - t e r m i n a l s p i n d l e attachm ent r e g io n . 43.
M a g n i f i c a t i o n x 13&0.
P h o t o m i c r o g r a p h o f d i p l o i d an d t e t r a p l o i d r e s t i n g c e l l s o f A l l i u m v a l l d u m t a k e n from an a c e t o - n i g r o s l n e sm e a r p r e p a r a t i o n . M a g n i f i c a t i o n x 14Q0.
L_
_J
70
F ig . 43
L
71 r
i
E x p la n a tio n of F ig u re s 44.
P h o t o m i c r o g r a p h a p a i r e d and r e l a t i o n a l l y c o i l e d e a r l y p r o p h a s e of A l l i u m v a l l d u m t a k e n fr o m an a c e t o - n i g r o s i n e paratio n .
45.
sm e ar p r e
M a g n ifIc e tio n x 1350.
P h o t o m i c r o g r a p h o f a p a i r e d and r e l a t i o n a l l y c o i l e d m id -prop hase of Allium v a lld u m taken from an a c e t o - n i g r o s i n e s m e a r p r e p a r a t i o n . M a g n i f i c a t i o n x 1400.
L
72
75 r
E xplanation of Figures 46.
P h o to m ic ro g ra p h of a p a i r e d and r e l a t i o n a l l y c o i l e d p o l y p l o i d prom etaphase of A llium v a l l d u m t a k e n fro m an a c e t o - n i g r o s i n e preparation.
47.
sm ear
M a g n i f ic a t i o n x 1550.
P h o t o m i c r o g r a p h o f a p o l y p l o i d m etap ha.se of A l l i u m v a l l d u m t a k e n from an a c e t o - n i g r o s i n e smear p r e p a r a t i o n sho wing a w id e s e p a r a t i o n of th e c h ro m a tid s .
l_
M a g n ific a tio n x 1550.
74 n
—
*
F i g . 46
F ig . 47
75
r
i
E x p la n a tio n of F ig u re s 48.
P ho to m icrog raph of a p o l y p lo i d anaphase of A l l i u m v a l i d u m t a k e n fro m an a c e t o - n i g r o s i n e sm ear p r e p a r a t i o n .
49.
M a g n i f i c a t i o n x 1350*
Photom icrog raph of a p o l y p lo i d m etaphase of A l l i u m v a l i d u m t a k e n f ro m an a c e t o - n i g r o s i n e sm ear p r e p a r a t i o n sh o w in g t h e c h r o m a t i n lum p. M ag n ificatio n x 1350.
I_
77 r
E x p lan atio n of Figure 50.
Photom icrograph of a l o n g i t u d i n a l s e c t i o n o f a y o u n g s e e d l i n g 15 mm. l o n g o f A l l i u m v a l l d u m s h ow in g t h e v a r i o u s t i s s u e s . M a g n ific a tio n x 175.
L
ECT =
ectoderm is
FL
=
first
RT
=
r e g i o n of t r a n s i t i o n
C
=
co tyledo n
CC
=
c o r t e x of c o t y l e d o n
CT
=
c o r t e x of r e g i o n o f t r a n s i t i o n
leaf
78 n
I
*
F i g . 50
79 r
E xplanation of Figures 51.
Photom icrograph of a b i n u c l e a t e c e l l of A l l i u m v a l l d u m t a k e n fro m an a c e t o - n i g r o s i n e sm ear p r e p a r a t i o n s h o w in g a l a r g e n u c l e u s and a m i c r o n u c l e u s .
52.
M a g n i f i c a t i o n x 1550.
P h o to m icro g rap h of a t e l o p h a s e of A lliu m v a l l d u m t a k e n from an a c e t o - n i g r o s i n e sm ear p r e p a r a t i o n s ho w ing t h e f o r m a t i o n o f a m icro n u cleu s.
L
M a g n i f ic a t i o n x 1550.
80 r~ n
F i g * 51
E x p la n a t i o n of F ig u re P hotom icrograph of a te lo p h a s e of A llium v a l l d u m sho wing b r i d g e s a n d f r a g m e n t s . An a c e t o - n i g r o s i n e sm ear p r e p a r a t i o n . M a g n ific a tio n x 1250.
82
F i g * 53
L
E xp lan atio n of Figure Photom icrograph of a dum b-bell shaped n u cleu s o f A l l i u m v a l l d u m t a k e n fro m a n a c e t o n i g r o s i n e smear p r e p a r a t i o n . x 1400.
M agnification
84
——
“i
F i g . 54
6
85
~i
the M editerranean, p a r t s of t h e w o r l d .
i t h a s b e e n n a t u r a l i z e d i n many o t h e r I t d o e s n o t fo rm a l a r g e ,
flatten ed
b u l b b u t h a s an e l o n g a t e d , c y l i n d r i c a l one w h i c h i s t h e edible p o rtio n .
The p l a n t h a s a ste m two t o t h r e e f e e t
h i g h w h i c h b e a r s a n umbel o f w h i t e f l o w e r s marked w i t h p u rp le. Levan ( 1 9 5 1 ) r e p o r t e d t h e d i p l o i d chromosome n um b er f o r A l l i u m p o rru m t o be t h i r t y - t w o .
Twenty-
e i g h t o f t h e chromosomes h a v e m ed ian o r s u b - m e d i a n sp in d le attachm ent re g io n s g iv in g r i s e
to the V-shaped
c o n f i g u r a t i o n common t o t h i s g e n u s ( f i g .
55).
Four
o f t h e chromosomes h a ve s u b - t e r m i n a l s p i n d l e a t t a c h m e n t regions.
It
i s t o t h e s h o r t e r arms o f t h e s e t h a t
r a th e r large s a t e l l i t e s
ere a ttac h ed ( f i g .
56).
P o l y p l o i d y was a r e g u l a r b u t i n f r e q u e n t f a c t o r i n c l u d e d i n t h e d e v e l o p m e n t o f t h e you ng s e e d l i n g s . T e t r a p l o i d y was a l w a y s f o u n d t o be p r e s e n t i n c e l l s t h a t were l a r g e r t h a n t h e d i p l o i d c e l l s
( f ig . 5?).
The chromosomes o f t h e p o l y p l o i d c e l l s r e m a i n e d p a i r e d and r e l a t i o n a l l y and 5 9 ) .
c o i l e d t h r o u g h o u t p r o p h a s e ( f i g s . 58
At m e t a p h a s e , t h e chromosomes s e p a r a t e d and
g a v e r i s e t o s i x t y - f o u r chromosomes ( f i g . 6 0 ) . d i v i s i o n went t o c o m p l e t i o n n o r m a l l y ( f i g . 6 1 ) .
The O cto-
p l o i d y was n e v e r f o u n d . By means of s e c t i o n s ,
t h e r e g i o n i n w h ic h t h i s
phenomenon o c c u r r e d was d e t e r m i n e d .
The c o r t e x o f t h e
c o t y l e d o n and of t h e t r a n s i t i o n r e g i o n b e t w e e n r o o t and shoot c o n ta in e d p o ly p lo id c e l l s
( f ig . 62),
A few p o l y
p l o i d c e l l s we re p r e s e n t i n t h e e c t o d e r m i s o f t h e cotyledon. The f i r s t e v i d e n c e o f t h e o c c u r r e n c e of p o l y p l o i d y was s e e n i n s m e a r s o f s e e d l i n g s 10 mm. l o n g .
I t was no
l o n g e r fo u n d a t 44 mm. a l t h o u g h some we re f o u n d t o be d e v o i d o f t h e s e c e l l s a t 38 mm..
T h i s was d i f f i c u l t t o
d e t e r m i n e s i n c e t h e p o l y p l o i d c e l l s w ere r a r e t h r o u g h out the e n tir e - ^ in v e s tig a tio n .
However, a t 44 mm., no
d i v i s i o n s of any k i n d w ere f o u n d i n t h e c o r t e x b u t d i p l o i d d i v i s i o n s were s t i l l e v i d e n t i n t h e p r l m o r d i u m of the f i r s t
leaf.
The f i r s t l e a f a p p e a r e d a t 65 t o 70 mm..
The a b e r r a t i o n s o b s e r v e d w ere s i m i l a r i n e v e r y r e s p e c t to th o se r e p o r te d p r e v io u s ly f o r the o th e r species.
The b i n u c l e a t e c e l l s c o n s i s t e d o f a l a r g e
n u c l e u s a nd a m i c r o n u c l e u s ( f i g . 6 3 ) . stage fre q u e n tly
i n c l u d e d b r i d g e s and chro m osom al
fragm ents ( f i g . 6 4 ). w ere a l s o f o u n d .
The t e l o p h a s e
Dumb-bell shaped n u c l e i ( f i g . 65)
None o f t h e s e a b n o r m a l i t i e s were
very f r e q u e n t. A l l i u m ammophllum H e u f f A l l i u m ammophllum. a member of t h e g a r l i c g r o u p , i s a n a tiv e of Europe. lavender flo w ers.
I t b e a r s l o o s e c l u s t e r s of
87 r
i
E xplan atio n of F ig u res 55.
Photom icrograph of a
d i p l o i d c e l l of Allium
p o rr u m t a k e n from a n
a c e t o - o r c e i n sm ear p r e
p a r a t i o n s ho w ing t h e c o n f i g u r a t i o n o f t h e c h ro m o s o m e s. 56.
M a g n i f ic a t i o n x 1400.
P h o to m icro g rap h of a p o rr u m t a k e n fro m a n
d i p l o i d c e l l of A llium a e e vt o - o r c e i n s m e a r p r e
p a r a t i o n show in g a l a r g e s a t e l l i t e t o t h e s h o r t e r arm o f a chro m osome. M a g n i f ic a t i o n x 1550.
L
attach ed
88 1
F i g . 55
F ig . 56 L
_l
89 r
i
E xplanation of F igures 57.
P h o t o m i c r o g r a p h o f a d i p l o i d c e l l and a t e t r a p l o i d c e l l o f A lliu m pdrrum t a k e n from an a c e t o - o r c e i n sm ear p r e p a r a t i o n .
M agnification
x 1350. 58.
P h o t o m i c r o g r a p h o f a p a i r e d and r e l a t i o n a l l y c o i l e d p o l y p l o i d p ro p h a s e o f A lliu m porrum t a k e n f r o m a n a c e t o - o r c e i n sm e ar p r e p a r a t i o n . M a g n if ic a tio n x 1350.
L_
90 n
F i g . 57
F ig . 58 L.
91 r
E x p la n a tio n of F ig u re s 59.
Photom icrograph of a p o ly p lo id m id-prophase A l l i u m p o r r u m t a k e n fro m an a c e t o - o r c e i n sm e ar p r e p a r a t i o n s h o w in g p a i r i n g and re la tio n a l co ilin g .
60.
M a g n if ic a tio n x 1550.
P hotom icrograph of a p o l y p lo i d m etaphase of A l l i u m p o r r u m sh o w in g s i x t y - f o u r chromosomes w ith w idely se p a ra te d c h ro m a tid s . o r c e i n sm ear p r e p a r a t i o n .
An a c e t o -
M agnification
x 1400.
_j
92
E xplan atio n of F ig u re s Photom icrograph of a p o ly p lo id anaphase of A l l i u m p o r r u m t a k e n f ro m an a c e t o - o r c e i n sm ear p r e p a r a t i o n .
M a g n if ic a tio n x 1400.
94 “ i
L
95 r
E x p lan atio n of Figure 62.
Photo m icrog raph of a l o n g i t u d i n a l s e c t i o n o f a y ou n g s e e d l i n g 25 mm. l o n g o f A l l i u m p o r r u m sh o w in g t h e v a r i o u s t i s s u e s . M a g n i f ic a t i o n x 175.
L_
EGT =
ectoderm is of co ty led o n
FL =
f i r s t leaf
RT =
region of t r a n s itio n
C
cotyledon
=
CG =
c o rte x of coty led on
CT =
c o r t e x of r e g io n of t r a n s i t i o n
9 6
n
* a
f V " 4 > I* I •
^
» t | *f
F i g . 62
; -# -2
9
?
r
E xplanation of F ig u res 63.
P h o to m icro g rap h of a b i n u c l e a t e c e l l of A llium p o rr u m t a k e n from an a c e t o - o r c e i n sm e a r p r e p a r a t i o n s h o w in g a l a r g e n u c l e u s and a m i c r o nucleus.
64.
M a g n i f i c a t i o n x 1350*
P h o to m icrog rap h of a t e l o p h a s e o f Allium p o r r u m t a k e n from a n a c e t o - n i g r o s i n e
smear
p r e p a r a t i o n sho w ing c hro m osom a l b r i d g e s and fragm ents.
L_
M a g n if ic a tio n x 1400,
_i
98
E x p lan atio n of Figure Photom icrograph of a dum b-bell shaped n u c le u s o f A llium porrum ta k e n from an a c e t o - o r c e i n sm ear p r e p a r a t i o n . M a g n if ic a tio n x 1400,
100 n
F i g . 65
L
101 r
-1 T h i s s p e c i e s h a s had two d i p l o i d chromosome numbers reported,
s i x t e e n and t h i r t y - t w o ( L e v a n , 1 9 3 5 ) .
The
p a r t i c u l a r one u s e d h e r e had a chromosome number o f th irty -tw o .
Previous in v e s tig a tio n s
( L e v a n , 1935) was
done on t h e one w i t h a d i p l o i d number o f s i x t e e n . th is case,
In
i t was r e p o r t e d t h a t a l l o f t h e chromosomes
h a d a m ed ian s p i n d l e a t t a c h m e n t r e g i o n and t h a t t h e r e was d o u b t c o n c e r n i n g t h e e x i s t e n c e o f s a t e l l i t e s . Although t h i s
i n v e s t i g a t i o n was n o t u n d e r t a k e n f o r t h e
d e term in atio n of karyotype,
i t may be s e e n from f i g u r e s
69 and 70 t h a t t h e chromosomes o f A l l i u m ammophilum h a v e m e d ia n and s u b - m e d i a n s p i n d l e a t t a c h e m e n t r e g i o n s . P o l y p l o i d y was f a r more a b u n d a n t i n t h i s s p e c i e s t h a n i n any of t h e o t h e r s s t u d i e d . and f r e q u e n t l y
I t occurred r e g u la r ly
i n t h e y o un g s e e d l i n g s .
The c e l l s i n
g e n e r a l were l a r g e b u t t h e t e t r a p l o i d c e l l s were a l w a y s g r e a t e r in d ia m e te r than the d i p lo i d c e l l s
( f i g . 66).
The chromosomes r e m a i n e d c l o s e l y p a i r e d and r e l a t i o n a l l y c o i l e d t h r o u g h o u t p r o p h a s e ( f i g s . 67 and 6 8 ) u n d e r g o i n g a s e p a r a t i o n a t m e t a p h a s e t o form s i x t y - f o u r chromosomes ( f i g s . 69 and 7 0 ) . A n o t h e r s e p a r a t i o n f o l l o w e d t h i s in which th e c h ro m a tid s s e p a r a t e d .
T h i s marked t h e
b e g i n n i n g o f a n a p h a s e w h i c h was n o r m a l ( f i g . the follo w in g te lo p h a s e .
O c t o p l o i d c e l l s were n o t s e e n .
P o l y p l o i d d i v i s i o n s were f i r s t s e e d l i n g s 10 mm. l o n g . L
.
7 1) a s was
o b s e r v e d i n young
These d i v i s i o n s we re v e r y -i
102 “i
num ero us u n t i l i t r e a c h e d t h e l e n g t h o f 57 mm.
From
h e r e on, t h e c o r t i c a l d i v i s i o n s d e c r e a s e d u n t i l t h e y d i s a p p e a r e d e n t i r e l y a t 43 mm.
At t h i s l e n g t h , no
d i v i s i o n s o f any k i n d were s e e n e x c e p t i n t h e m e r i s t e m of th e f i r s t
leaf.
The f i r s t
l e a f was v i s i b l e m a c r o -
s c o p i c a l l y a t 65 t o 75 mm.. L o n g i t u d i n a l s e c t i o n s of t h e young s e e d l i n g showed t e t r a p l o l d c e l l s t o be p r e s e n t i n t h e c o r t e x of b o t h t h e c o t y l e d o n and r e g i o n o f t r a n s i t i o n b e t w e e n r o o t and shoot.
O c c a s i o n a l l y p o l y p l o i d c e l l s were s e e n i n t h e
ectoderm is ( f i g . 72). A l l i u m ammoph1 1 urn d i d n o t p r e s e n t e v i d e n c e f o r any a b n o r m a l i t i e s o t h e r t h a n t h o s e r e p o r t e d f o r t h e other species.
D u m b - b e l l shap ed n u c l e i ,
and f r a g m e n t s ( f i g .
telophase bridges
7 3 ) , and b i n u c l e a t e c e l l s w i t h a
m a c r o n u c l e u s and a m i c r o n u c l e u s ( f i g . 7 4 ) were s e e n . The d u m b - b e l l s h a p e d n u c l e i and t e l o p h a s e f r a g m e n t s w e re r a r e b u t t h e b r i d g e s and m i c r o n u c l e i were n u m e r o u s . A l l i u m t u b e r o s u m L. A l l i u m t u b e r o s u m , a l s o known a s C h i n e s e C h i v e s , indigenous to the Far E a s t.
The b u l b s a r e c l u s t e r e d and
n a r r o w b e i n g s c a r c e l y one c e n t i m e t e r t h i c k .
The umbel
b e a rs w hite s t e l l a t e flo w e rs . The d i p l o i d chromosome number i s t h i r t y - t w o (La C o u r , u n p u b l . ) .
W ith t h e e x c e p t i o n o f t h i s w ork ,
is
103 r
'i
E x p la n a tio n of F ig u r e s 66.
P h o t o m i c r o g r a p h of a d i p l o i d n u c l e u s a nd a t e t r a p l o i d n u c l e u s o f A l l i u m ammophllum t a k e n fro m an a c e t o - n i g r o s i n e sm e a r p r e paratio n .
67.
M a g n ific a tio n x 1350.
Photom icrograph of a p o ly p lo id e a r l y p r o p h a s e o f A l l i u m ammophllum t a k e n f r o m an a c e t o - n i g r o s i n e sm e a r p r e p a r a t i o n s h o w i n g p a i r i n g and r e l a t i o n a l c o i l i n g . M a g n i f i c a t i o n x 1 400.
L
104
Fig* 67 L
105 r
E x p la n a tio n of F ig u r e s 68.
Photom icrograph of a p o ly p lo id m id-prophase A l l i u m ammophilum t a k e n from an a c e t o nigrosine
sm ear p r e p a r a t i o n and sh o w in g
r e l a t i o n a l c o l l i n g and p a i r i n g .
M agni
f i c a t i o n x 1400. 69.
P ho to m icro grap h of a p o ly p lo i d m etaphase of A l l i u m ammophllum t a k e n fro m a n a c e t o n i g r o s i n e sm e a r p r e p a r a t i o n and show in g s i x t y - f o u r c h ro m o s o m e s. x 1 35 0 .
L_
M agnification
106
F ig * 68
F ig . 69
107 r
E x p la n a tio n of F ig u re s 70.
P hotom icrograph of a p o l y p lo i d m etaphase o f A l l i u m ammophilum t a k e n fr om a n a c e t o nigro sine
sm ear p r e p a r a t i o n and sho w in g
s i x t y - f o u r g r e a t l y c o n t r a c t e d c h ro m o s o m e s . M a g n ific a tio n x 1250. 71.
Pho tom icro grap h of a polyploid, anaphase of A l l i u m ammophilum t a k e n from an a c e t o n i g r o s i n e sm e a r p r e p a r a t i o n . x 1250.
L
M agnification
108
—
f
F ig.
?Q
F ig . 71 L -J
109 r
i
E x p l a n a t i o n of F i g u r e 72.
P h otom icro grap h of a l o n g i t u d i n a l s e c t i o n o f a s e e d l i n g o f A l l i u m ammophllum 18 mm. l o n g s h o w in g t h e v a r i o u s t i s s u e s . M a g n i f ic a t i o n x 175.
L.
EOT =
ecto d erm is of c o ty led o n
FL
first
= f
leaf
*
RT
=
r e g i o n of t r a n s i t i o n
C
=
cotyledon
cc
=
c o r t e x of c o ty l e d o n
OT
=
c o r t e x of r e g i o n o f t r a n s i t i o n
110 “I
* I
~ *
** * * * v* « to%-■ w * ♦
.
t
*”
%
*
ft
•
*
* * —s------ cc **•
jh
• •
- • *
4?
** *
* *
.
•
*- - - - - - - ECT
IS ! i
•
. ‘ •*% • *
* •
#
#
•* #
#|
* * % V '* ♦ * * 4
'
• , *
5 ■ ■"
#
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*
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*
■ ’%
*- - - - - - - - - - - - - - - - - - - - - - - - - R T \
%
*•
.
*
i * .* • ? - * [ '
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•
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-
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ii.ili. ** v -Mel*ft
Fig* 72
#
,9 ;
I ll r
'i
E x p lan atio n of F ig u res 75.
P h o t o m i c r o g r a p h o f a d i p l o i d t e l o p h a s e of A l l i u m ammophllum t a k e n fro m an a c e t o nlgrosine
sm e ar p r e p a r a t i o n a nd s h o w in g
c hro m osom a l b r i d g e s and f r a g m e n t s . M a g n ific a tio n x 1550. 74.
Photom icrograph of a b i n u c l e a t e c e l l of A l l i u m ammophllum t a k e n fro m an a c e t o n i g r o s i n e s m e a r p r e p a r a t i o n and s h o w in g a l a r g e n u c l e u s and a m i c r o n u c l e u s . M a g n ific a tio n x 1550,
L.
_ j
112
L
little
e l s e h a s b e e n done on t h i s s p e c i e s .
The c h ro m o
somes a p p e a r t o h a v e m e d i a n o r s u b - m e d i a n s p i n d l e a t t a c h ment r e g i o n s ( f i g .
74),
I t was n o t d e t e r m i n e d w h e t h e r
s a t e l l i t e s w e re p r e s e n t . I t was f o u n d t h a t t h e young s e e d l i n g s of A l l i u m t u b e r o s u m r e g u l a r l y and f r e q u e n t l y f o r m e d t e t r a p l o i d c ells.
These c e l l s u n d e r w e n t n o r m a l m i t o s i s .
p lo id d iv is io n s occurred in la rg e c e l l s
(fig .
The p o l y 76).
The
chromosom es w e re a l w a y s s e e n t o be c l o s e l y p a i r e d and r e l a t i o n a l l y c o i l e d th ro u g h p ro p h ase t o metaphase ( f i g s . 7 7 , 7 8 , 79 , and 8 0 ) .
The s p i n d l e a t t a c h m e n t r e g i o n
o f t h e s e chromosomes r e m a i n e d s i n g l e and u n d i v i d e d u n t i l v e r y l a t e p r o p h a s e o r e a r l y m e t a p h a s e ( f i g s . 79 a nd 8 0 ) . At m e t a p h a s e t h e p a i r e d chromosomes s e p a r a t e d .
T his
was f o l l o w e d by a n o t h e r s e p a r a t i o n w h i c h i n i t i a t e d a n a p h a s e movement w h i c h was n o r m a l ( f i g . 8 1 ) .
No o c t o -
p l o i d d i v i s i o n s w ere o b s e r v e d . The c o r t e x o f b o t h t h e c o t y l e d o n and r e g i o n o f t r a n s i t i o n w ere f o u n d t o be t h e r e g i o n s of t h e young s e e d l i n g s i n w h i c h m ost o f t h e p o l y p l o i d c e l l s o c c u r r e d . A few t e t r a p l o i d c e l l s we re p r e s e n t i n t h e e c t o d e r m l s (fig . 82).
The l o w e r p o r t i o n o f t h e c o t y l e d o n c o n t a i n e d
more p o l y p l o i d c e l l s t h a n t h a t r e g i o n c l o s e r t o t h e s e e d When t h e s e e d l i n g s w ere 10 mm. l o n g , p o l y p l o i d d i v i s i o n s we re f i r s t
observed.
This c o n d it i o n p e r s i s t e d
u n t i l t h e s e e d l i n g s m e a s u r e d 40 mm.. At t h i s p o i n t ,
114
no d i v i s i o n s , in the co rtex*
e i t h e r d i p l o i d o r t e t r a p l o i d were f o u n d Some d i v i s i o n s w ere p r e s e n t i n t h e
m eristem a tic re g io n of the f i r s t a l l d ip lo id .
The f i r s t
lo n g itu d in al s l i t
3e a f b u t t h e s e we re
l e a f appeared th rough the
i n t h e c o t y l e d o n o u s s h e a t h a t 65 t o 75
mm, . In a d d i t i o n t o th e p o l y p l o i d c e l l s ,
the abnor
m a l i t i e s r e p o r t e d f o r a l l th e o th e r s p e c i e s used i n t h i s i n v e s t i g a t i o n w ere f o u n d t o e x i s t i n t h e same r e g i o n w ith the t e t r a p l o i d c e l l s .
B inucleate c e l l s w ith a
m a c r o n u c l e u s and a m i c r o n u c l e u s ( f i g . 8 3 ) w e re p r e s e n t . D u m b - b e l l s h a p e d n u c l e i ( f i g s . 84 and 8 5 ) and t e l o p h a s e b r i d g e s and f r a g m e n t s ( f i g . 8 6 ) were f o u n d .
115 r
E x p la n a tio n of F ig u re s 76.
P h o t o m i c r o g r a p h of a d i p l o i d m e t a p h a s e o f A l l i u m t u b e r o s u m show in g t h e c o n f i g u r a t i o n an d w i d e l y s p r e a d c h r o m a t i d s . o r c e i n sm e ar p r e p a r a t i o n . X
76.
An a c e t o -
M agnification
1400*
P h o t o m i c r o g r a p h o f a d i p l o i d c e l l and a t e t r a p l o i d c e l l of A l l i u m t u b e r o s u m t a k e n fr o m an a c e t o - n i g r o s i n e M a g n i f ic a t i o n x 1550.
L
sm e ar p r e p a r a t i o n *
116
F ig . 76
L
11? [~
E x p la n a tio n of F ig u re s 7?.
P h o t o m i c r o g r a p h o f an e a r l y p o l y p l o i d p r o p h a s e o f A l l i u m t u b e r o s u m t a k e n f r o m an a c e t o - n l g r o s i n e s m e a r p r e p a r a t i o n and show in g p a i r i n g and r e l a t i o n a l c o i l i n g . M ag n ificatio n x 1400.
78.
P h o t o m i c r o g r a p h of a l a t e p o l y p l o i d p r o phase of A lliu m tuberosum t a k e n from an a c e t o n i g r o s i n e s m e a r p r e p a r a t i o n and s h o w in g r e l a t i o n a l c o i l i n g and p a i r i n g . t i o n x 1400.
L_
M agnifica
I;.'
118 n
F ig . 78
119 r
E x p la n atio n of F ig u res 79.
P h o to m ic ro g ra p h of a p o l y p l o i d m etaphase A l l i u m t u b e r o s u m s h o w in g t e t r a c h r o m o somes. tio n .
80.
An a c e t o - n i g r o s i n e s m e a r p r e p a r a M a g n ific a tio n x 1400.
P h o to m icro g rap h of a p o ly p lo id m etaphase A l l i u m t u b e r o s u m i n w h i c h some o f t h e s p in d le a tta c h m e n t r e g i o n s have s e p a r a t e d . An a c e t o - n i g r o s i n e sm e a r p r e p a r a t i o n . M agnification x 1350.
120
E x p la n atio n of Figure P ho to m icro g rap h of a p o l y p lo i d an aph ase A l l i u m t u b e r o s u m t a k e n fr o m a n a c e t o n i g r o s i n e smear p r e p a r a t i o n . f i c a t i o n x 1550.
M agni
1 22
E xp lanatio n of Figure P h o t o m i c r o g r a p h of a l o n g i t u d i n a l s e c t i o n o f a y o un g s e e d l i n g 20 mm. l o n g o f A l l i u m t u b e r o s u m sh o w in g t h e v a r i o u s t i s s u e s . M a g n if ic a tio n x 175, EOT
ecto d erm is of c o ty le d o n
FL
f i r s t leaf
RT
r e g io n of t r a n s i t i o n
C
cotyledon
CO
c o rte x of cotyledon
CT
c o r t e x of r e g i o n of t r a n s i t i o n
124 n
cc ECT
FL
CT -RT F i g * 82
125 r
i
E xp lan atio n of Figures 83.
P h o to m icro g rap h of a b i n u c l e a t e
A llium
t u b e r o s u m t a k e n f ro m an a c e t o - n i g r o s i n e
sm ear
p r e p a r a t i o n and sho w in g a l a r g e
and a
m icronucleus. 84.
c e l l of
nucleus
M a g n ific a tio n x 1550.
P h o t o m i c r o g r a p h o f two d a u g h t e r c e l l s o f Allium tuberosum r e v e r t i n g t o th e r e s t i n g con d itio n .
The ch rom oso m a l b r i d g e r e m a i n s u n
b r o k e n b e t w e e n them a nd f o r m s a t y p e o f dumb b e l l shaped n u c le u s . preparation.
L
An a c e t o - n i g r o s i n e
M a g n i f ic a t i o n X 1350,
sm ear
126
F ig , 84 L
127 r
E x p la n a tio n of F ig u r e s 85*
P h o to m icro g rap h of a du m b -b ell shaped n u c le u s of* A l l i u m t u b e r o s u m t a k e n fro m an a c e t o n i g r o s i n e smear p r e p a r a t i o n .
M agnification
x 1550. 86.
P h otom icrograph of a t e l o p h a s e of A llium t u b e r o s u m t a k e n fro m an a c e t o - n i g r o s i n e sm ear p r e p a r a t i o n a n d s ho w ing chromosome b r i d g e s and f r a g m e n t s .
L_
M a g n i f i c a t i o n x 1550.
128 n
>
F i g . 35
Ky*x
*
t m
F ig . 86 _1
L
129 r
~i
I V . DISCUSSION Stomps ( 1 9 1 0 ) an d W i n k l e r ( 1 9 1 6 ) f i r s t c o u n t e d t h e number o f chromosomes i n p o l y p l o i d c e l l s
of d i p l o i d p l a n t s .
D i r e c t e v i d e n c e f o r t h e p r e s e n c e o f p o l y p l o i d y was a l s o a d v a n c e d by G e i t l e r ( 1 9 3 9 ) .
In h i s i n v e s t i g a t i o n s of
G e r r i s , he n o t i c e d t h a t t h e s e x chromosome r e m a i n e d c o n d e n s e d and h e t e r o p y c n o t i c i n t h e r e s t i n g n u c l e u s . a u t o s o m e s became r e t i c u l a t e . was p o s s i b l e
The
By means of t h i s f a c t o r ,
it
t o d e t e r m i n e t h e d e g r e e of p o l y p l o i d y by
c o u n t i n g t h e number of x chromosomes p r e s e n t i n e a c h c e l l . P r i o r t o t h e s e c a s e s , most e v i d e n c e had b e e n o f an i n d i r e c t n a t u r e s u c h a s t h e s i z e of t h e r e s t i n g c e l l (Meurman, 1 9 3 3 ) .
T h i s c o u l d l e a d t o e r r o r due t o
v a c u o l i z a t i o n and c e l l s p e c i a l i z a t i o n . N uclear s i z e has a l s o been used as a c r i t e r i o n a nd i s p r o b a b l y of more v a l u e t h a n c e l l s i z e s i n c e t h e y are l e s s su b je ct to v a r i a t i o n , increase
G e i t l e r ( 1 9 4 1 ) s a i d an
i n n u c l e a r s i z e may o c c u r b e c a u s e of i n n e r
m u l t i p l i c a t i o n of chromosomes or an i n c r e a s e i n t h e s i z e o f t h e i n d i v i d u a l chromosome when t h e d i p l o i d number I s m a i n t a i n e d .
The l a t t e r p o s s i b i l i t y
therefore
would c a u s e an e n l a r g e m e n t o f t h e n u c l e u s w i t h o u t t h e presence of p o ly p lo id y th u s g iv in g r i s e th is
to e r r o r i f
method a l o n e were u s e d a s th e d e t e r m i n i n g f a c t o r . De Mol ( 1 9 2 7 ) c o u n t e d t h e number o f n u c l e o l i i n
L.
re stin g n u clei.
N u c l e o l i , however, have s ten d en cy to
f u s e th u s r e d u c i n g t h e number.
The maximum number o f
n u c l e o l i a re found in t e l o p h a s e .
C o n s e q u e n t l y some h a v e
p r o b a b l y f u s e d by r e s t i n g s t a g e . B e r g e r ( 1 9 4 1 ) f o u n d a new d i r e c t means o f d e t e r m ining th e d e g re e of p o l y p lo i d y i n r e s t i n g n u c l e i of S oinacia. lite s
T h i s was t h e number o f h e t e r o p y c n o t i c s a t e l
e i t h e r i n d i r e c t c o n t a c t w ith th e n u c le o lu s or
j o i n e d t o i t by a s h o r t t h r e a d .
He s a y s , h o w e v e r ,
that
t h i s method would be d i f f i c u l t and I m p r a c t i c a b l e w i t h more t h a n one p a i r o f s a t e l l i t e d c h ro m o s o m e s.
D irect
chromosome c o u n t s seemed most s u i t a b l e f o r t h i s in v estig atio n . Ever s in c e th e d is c o v e ry of p o ly p lo id y (W inkler, 1 9 1 6 ) , many t h e o r i e s h ave b e e n p r o p o u n d e d t o e x p l a i n t h i s phe nomenon.
W i n k l e r ( 1 9 1 6 ) p r o p o s e d two ways i n
w h i c h s u c h c e l l 6 may h a v e o r i g i n a t e d .
He s t a t e d t h a t
i t may be due t o t h e f u s i o n o f two n u c l e i . h a s a l s o b e e n h e l d by B r e z l a v e t z ( 1 9 5 2 ) .
T his th e o ry T his s i t u a t i o n
c a n n o t o b t a i n i n th e s p e c i e s of A llium i n v e s t i g a t e d b e c a u s e o n l y p a i r e d p o l y p l o i d p r o p h a s e s were o b s e r v e d . If
two n u c l e i h a d f u s e d , a n u n p a i r e d c o n d i t i o n would
e x i s t and r e l a t i o n a l c o i l i n g would n o t be p r e s e n t . Such a c o n d i t i o n was n e v e r f o u n d .
The same h o l d s t r u e
f o r h i s second e x p l a n a t i o n t h a t the b i n u c l e a t e c e l l s w h i c h a r e a l r e a d y p r e s e n t i n stem t i s s u e s u n d e r g o
m i t o s i s and r e v e a l t h e m s e l v e s t o be p o l y p l o i d .
Again,
t h i s would g i v e r i s e
to u n p a ire d p ro p h ases la c k in g
re la tio n a l co ilin g .
T h i s s t a t e was n o t o b s e r v e d .
Nemec ( 1 9 3 1 ) s u g g e s t e d t h a t t e t r a p l o i d c e l l s may a r i s e th ro u g h a second l o n g i t u d i n a l s p l i t m e t a p h a s e chro m osom e.
in each
T h i s would mean t h a t t h e m e t a p h a s e
p l a t e w o u ld f i r s t h a v e t h e d i p l o i d number o f c h rom o so m e s, e a c h chromosome h a v i n g two c h r o m a t i d s .
These would t h e n
undergo a l o n g i t u d i n a l s e p a r a t i o n form ing m etaphase chromos omes w i t h a s i n g l e
chrom atid.
As may be s e e n i n
f i g u r e s 4 7 , 6 0 , 7 0, and 7 5, t h e m e t a p h a s e chromosome c o n s i s t s of two c h r o m a t i d s .
A s e p a r a t i o n , however,
d o e s o c c u r b u t t h i s m arks t h e b e g i n n i n g o f a n a p h a s e . The m e t a p h a s e chromosome i s
a lw a y s d o u b l e .
C o n sid e rin g th e c l o s e l y p a ir e d c o n d i t i o n of the chromos omes i n t h e p o l y p l o i d c e l l s ,
the r e l a t i o n a l
c o i l i n g and t h e u n d i v i d e d s p i n d l e a t t a c h m e n t r e g i o n s , i t w ould seem t h a t t h e s e chromosomes h a v e n o t u n d e r gone a d i v i s i o n s i n c e t h e y r e d u p l i c a t e t h e m s e l v e s . They m u s t , t h e r e f o r e ,
have u n d e r g o n e a d o u b l e r e d u p l i
c a t i o n in the p re c e e d in g r e s t i n g s t a g e .
I f the
c o n d i t i o n s o f r e l a t i o n a l c o i l i n g an d c l o s e p a i r i n g a r e f o u n d i n c e l l s w h i c h a r e known t o h a v e u n d e r g o n e a t l e a s t one m i t o s i s s i n c e t h e o n s e t o f g e r m i n a t i o n ,
th is
w ould p r e s e n t e v i d e n c e t h a t t h e d o u b l i n g h a d t a k e n p la c e sin c e the beginning of germ in atio n (B erger,
1941).
132 r~
A l b l z z l a ( B e r g e r and W l t k u s , 1 9 4 9 ) , p o l y p l o i d d i v i s i o n s a r e f o u n d j u s t a s so o n a s d i v i s i o n s a r e fo u n d * Not a l l o f t h e c e l l s a r e p a i r e d *
T h e r e f o r e some o f
them m ust have d i v i d e d i n t h e e m b ry o lo g y o f t h e seed* T his i s n o t th e c a se in th e A llium s p e c i e s i n v e s t i g a t e d . P o l y p l o i d d i v i s i o n s w e r e f o u n d a s e a r l y a s 8 mm.. The t y p e of p o l y p l o i d y d i s c u s s e d h e r e d i f f e r s g r e a t l y from p o l y s o m a t y and e n d o m i t o s i s .
The f o r m e r i s
s i m i l a r i n t h e s i n g l e f a c t t h a t I t o r i g i n a t e d b y ’a d o u b l e r e d u p l i c a t i o n o f t h e chromosomes i n t h e p r e c e e d l n g r e s t i n g s t a g e ( G e n t c h e f f and G u s t a f s s o n , 1939; B e r g e r , 1941).
I t d i f f e r s by i n c l u d i n g n o t o n l y t e t r a p l o i d
c e lls but a lso o ctap lo id c e l l s .
B o th a p a i r e d and an
u n p a i r e d c o n d i t i o n were o b s e r v e d i n d i c a t i n g t h a t t h e cells
underw ent a d i v i s i o n a f t e r r e d u p l i c a t i o n .
Since
o n l y p a i r e d p o l y p l o i d c e l l s w ere f o u n d i n A l l i u m , t h e y c o u l d n o t h a ve d i v i d e d s i n c e beco m in g t e t r a p l o i d 4, 14,
27, 3 5 , 4 6 , 6 8 , and 7 7 ) .
m ito sis,
also gives r i s e
an i n t r a n u c l e a r m i t o s i s .
(fig s.
The l a t t e r c a s e , e n d o
to p o ly p lo id c e l l s .
T his i s
No s p i n d l e i s f o r m e d .
The
chromos omes p a s s t h r o u g h s t a g e s G e i t l e r c a l l s e n d o p r o p h a s e , e n d o m e t a p h a s e and e n d o a n a p h a . s e . spindle is lack in g ,
Since a
t h e chromosomes do n o t u n d e r g o
p o l a r movement a t a n a p h a s e .
They s e p a r a t e s l i g h t l y
~and t h e n r e v e r t t o a r e s t i n g s t a g e c o n d i t i o n , c e l l s become t e t r a p l o i d
from e n d o a n a p h a s e .
The
The
155 r
1
stages c h a r a c te r is tic in A lliu m .
of t h i s p ro cess are not p re s e n t
T his e l i m i n a t e s th e p o s s i b i l i t y o f th e
p o ly p lo id c o n d itio n a r i s i n g through e n d o m lto sls. I n Mimosa p u d i c a ( W i t k u s and B e r g e r ,
1947),
it
was f o u n d t h a t c e r t a i n c e l l s i n t h e y o ung r o o t became t e t r a p l o i d by t h e same mechan ism as t h a t d e s c r i b e d f o r A llium .
These p o l y p l o i d c e l l s d i v i d e d o n l y once a s
p o ly p lo id s.
In b o th of th ese p l a n t s ,
then,
the t e t r a
p l o i d c e l l s d i v i d e o n l y once b e f o r e t h e y u n d e r g o d i f f e r e n t i a t i o n which p r e v e n t s f u r t h e r m i t o s i s . ( B e r g e r and W i t k u s , 1 9 4 9 ) ,
A lblzzla
on t h e o t h e r h a n d , h a s u n
p a i r e d t e t r a p l o i d d i v i s i o n s a s commonly a s p a i r e d .
At
tim e s t h e s e p a i r e d d i v i s i o n s a re very c l o s e l y p a i r e d , a t o t h e r tim es th e a s s o c i a t i o n i s n o t as i n t i m a t e .
Thus
i t d i f f e r s from Mimosa p u d i c a . e n d o m l t o s l s i n S p i n a c l a o l e r a c e a and t h e c a s e h e r e d i s c u s s e d . I n th e genus A l l i u m , t h e r e oc cu r i n n a t u r e forms w i t h d i f f e r e n t chromosome n u m b e r s .
D i p l o i d chromosome
n u m b e r s o f 1 4 , 16 , 28 and 52 a r e i n c l u d e d i n t h i s in v estig atio n . Involved, series.
It
i s c l e a r t h a t t h e r e a r e two s e r i e s
t h e ? -chromosome s e r i e s and t h e 8-chrom oso m e I n b o t h o f them , t h e r e o c c u r s a d i p l o i d and
a t e t r a p l o i d form . A theory to e x p la in the d i v e r s i t y
of th e b a s ic
number w i t h i n t h e g e n e r a i s b a s e d on t h e a s s u m p t i o n t h a t l o n g - a r m e d chrom osomes b r e a k , e a c h g i v i n g r i s e L
t o two e u b - t e r m l n a l l y c o n s t r i c t e d chrom osomes ( L e v a n , 1932).
If,
f o r e x a m p l e , one o f t h e chromosomes o f
A llium s t e l l a t u m broke,
i n w h i c h t h e d i p l o i d number i s f o u r t e e n ,
t h e r e s u l t would be s i x t e e n ch ro m o so m es,
two o f
them h a v i n g a s u b - t e r m i n a l s p i n d l e a t t a c h m e n t r e g i o n . This s i t u a t i o n e x i s t s d i p l o i d , number i s
i n A l l i u m f l s t u l o s u m i n w h ic h t h e
sixteen ( f ig .
12).
T h e r e a r e s e v e r a l f a c t o r s t h a t may a c c o u n t f o r the t e t r a p l o i d form s.
G i l e s (1939) red u c ed the c e l l u l a r
w ater d u rin g m ic ro sp o ro g e n esis in T r a d e s c a n t la .
He
found t h a t th e c o o r d i n a t i o n of t h r e e a p p a r e n t l y in d e p e n d e n t though n orm ally sy n c h ro n ize d e v e n ts ,
chromosome,
n u c l e a r and c e l l d i v i s i o n was d i s t u r b e d .
T his caused
t h e p r o d u c t i o n o f p o l y p l o i d m i c r o s p o r e s and p o l l e n grain s.
V a r i a t i o n s o f t h e w a t e r s u p p l y o f p l a n t s may
be an i m p o r t a n t f a c t o r i n t h e p r o d u c t i o n o f p o l y p l o i d g a m e t e s and c o n s e q u e n t l y o f p o l y p l o i d r a c e s . S t o c k w e l l ( 1 9 3 5 ) f o u n d p o l y p l o i d y t o be a common fe a tu r e in the C p u n tlas.
S e v e r e c o n d i t i o n s a nd wide
g e o g ra p h ic a l d i s t r i b u t i o n produced p l a n t s w ith h ig h er chromosome n u m b e r s .
Many members o f t h e g e n u s A l l i u m
had t h e i r o r i g i n i n one c l i m a t e and w e re n a t u r a l ! z e d i n many o t h e r s . have a r i s e n .
I n t h i s w a s, p o l y p l o i d s p e c i e s may
Sax ( 1 9 3 ? ) f u r t h e r e d e v i d e n c e f o r t h i s
f a c t by s u b j e c t i n g p l a n t s o f T r a d e s c a n t l a p a l u d o s a t o t e m p e r a t u r e c h a n g e s and f o u n d t h a t t h i s may c a u s e
135 r~
-i
chromosome d i v i s i o n s w i t h o u t n u c l e a r d i v i s i o n and c o n s e q uently p o ly p lo id y . The a b e r r a t i o n s w h i c h o c c u r r e d i n t h e same r e g i o n s a s t h e t e t r a p l o l d y p o r b a b l y h a v e no c o n n e c t i o n w i t h th e m . I t h a s b e e n shown by N i c h o l s ( 1 9 4 1 ) t h a t t e l o p h a s e b r i d g e s and f r a g m e n t s a r e of n o r m a l o c c u r r e n c e i n t h e r o o t s of allium s e e d lin g s .
A c c o r d i n g t o B e r g e r and W i t k u s ( 1 9 4 6 ) ,
t h e s e c e l l s a r e v e ry l i k e l y e l i m i n a t e d as developm ent continues*
I t is po ssib le
t h a t they f u r n i s h a s e x p la n a
t i o n f o r t h e o t h e r two a b n o r m a l i t i e s f o u n d .
The dumb
b e l l s h a p e d n u c l e i may be f o r m e d from u n b r o k e n b r i d g e s . T h i s i s e v i d e n t i n f i g u r e s 54 and 8 4 . b r o k e away a t b o t h e n d s , loose.
In th is case,
nucleus ( f i g . 8 3 ). lacking ,
I f these b rid g es
t h e b r i d g e would be l y i n g
i t r o u n d s up and fo r m s a m i c r o
When ch ro m o som al f r a g m e n t s a r e
o n l y one m i c r o n u c l e u s i s f o r m e d .
are p r e s e n t,
two m i c r o n u c l e i may f o r m , one f r o m t h e
b r i d g e s and one fr om t h e f r a g m e n t ( f i g s . T here i s a l s o t h e p o s s i b i l i t y
it
10 and 5 2 ) .
t h a t t h e s e two may f u s e
and f o r m a s i n g l e m i c r o n u c l e u s . d istin c t,
When t h e y
I n s te a d of rem aining
is p o s s ib le th a t these m icro n u clei a tta c h e d
t h e m s e l v e s t o t h e end o f t h e m a c r o n u c l e u s f o r m i n g a type of du m b -b ell shaped n u c le u s ( f i g s .
20 and 4 1 )
frequenter observed in the m a t e r i a l s t u d i e d . The s p e c i e s i n v e s t i g a t e d have t h e same g e n e r a l stru ctu re L
a s t h a t o f A l l i u m c e p a (Hayw ard, 193 8) and J
follow a s i m i l a r developm ental p a t t e r n . le n g th of the
How ever, t h e
s e e d l i n g a t which p o l y p lo i d d i v i s i o n s
occurred v a rie d w ith the s p e c ie s , d if fe re n c e being c o n s id e ra b le .
i n some c a s e s t h e
This i s v e ry l i k e l y
c o n n e c t e d w i t h t h e f o r m a t i o n of t h e f i r s t l e a f
since
the c o ty le c o n o u s l e a f d i e s s h o r t l y a f t e r th e ap p earan ce of th e f i r s t l e a f .
In A llium s t e l l a t u m . f o r i n s t a n c e ,
t h e f i r s t l e a f e m e r g e s from t h e s m a l l o p e n i n g i n t h e c o t y l e d o n o u s l e a f a t 20 t o 25 mm.. we re no l o n g e r f o u n d a t 25 mm..
Polyploid d iv is io n s
The f i r s t
l e a f of
A l l i u m s c h o e n o p r a s u m a p p e a r e d a t a p p r o x i m a t e l y 40 mm. w h i l e no p o l y p l o i d d i v i s i o n s were s e e n a t 53 mm..
An
o t h e r s p e c i e s w i t h s i x t e e n ch rom oso m es, A l l i u m f i s t u l o s u m was no l o n g e r o b s e r v e d t o I n c l u d e p o l y p l o i d c e l l s i n t h e c o r t e x o f t h e c o t y l e d o n o r t r a n s i t i o n when t h e s e e d l i n g m e a s u r e d 40 mm.. a t 50 t o 6 0 mm..
l e a f appeared
I n A l l i u m v a l l d u m . no p o l y p l o i d
d i v i s i o n s w ere s e e n a t 45 mm.. through the l a t e r a l and 70 mm..
The f i r s t
slit
The f i r s t
l e a f e m erged
i n t h e c o t y l e d o n b e t w e e n 60
Two of t h e t h r e e s p e c i e s w i t h t h i r t y - t w o
ch ro m o so m es, A l l i u m p o r r u m and A l l i u m t u b e r o s u m had sim ilar p a tte rn s ,
th at is ,
th e appearance of the f i r s t
l e a f a t 65 t o 75 mm. a n d t h e l a c k o f p o l y p l o i d y a t 40 t o 45 mm..
A l l i u m ammophllum was d e v o i d o f p o l y p l o i d
d i v i s i o n s a t 43 mm. and gave m a c r o s c o p i c e v i d e n c e o f the f i r s t
l e a f b e t w e e n 70 and 80 mm..
From t h i s
13?
e v i d e n c e , i t would seem t h a t t h e r e i s a d e f i n i t e r e l a t i o n b e t w e e n t h e l i m i t a t w h i c h p o l y p l o i d y o c c u r s and the fo rm a tio n of the f i r s t l e a f .
L
138 ~i
V. SUMMARY 1.
Some t e t r a p l o i d c e l l s a r e f o r m ed d u r i n g t h e n o r m a l d e v e l o p m e n t o f y o ung s e e d l i n g s o f A l l i u m s t e l l a t u m . A lliu m f i s t u l o s u m , A llium sc h o en o p ra su m , A llium an g u lo su m . A llium v a li d u m , A llium p o rru m , Allium ammophllum. and A l l i u m t ub fero sum .
2.
T h ese p o l y p l o i d c e l l s u n d e r w e n t n o r m a l m i t o s i s ,
3.
The chromosomes r e m a i n e d c l o s e l y p a i r e d and r e l a t i o n a l l y c o ile d throughout prophase,
4.
The s p i n d l e a t t a c h m e n t r e g i o n s r e m a i n e d s i n g l e and un d iv id ed u n t i l prom etaphase or m etaphase,
5.
The p a i r e d chromosomes s e p a r a t e d a t m e t a p h a s e f o r m i n g a t e t r a p l o i d number o f m e t a p h a s e c h ro m o s o m e s.
6.
The m e t a p h a s e s e p a r a t i o n was f o l l o w e d by a n o t h e r s e p a r a t i o n w h i c h marked t h e b e g i n n i n g o f a n a p h a s e .
7.
The t e t r a p l o i d c o n d i t i o n r e s u l t s f r o m a d o u b l e chromosome r e d u p l i c a t i o n i n t h e p r e c e e d l n g r e s t i n g stage.
8.
No o c t o p l o l d d i v i s i o n s w e re o b s e r v e d t o be p r e s e n t .
9.
W it h t h e e x c e p t i o n o f A l l i u m a n g u l o s u m . t h e t e t r a p l o i d c e i l s w e re f o u n d m a i n l y i n t h e b a s a l p o r t i o n of the c o ty le d o n .
10.
P o l y p l o i d d i v i s i o n s were f o u n d i n t h e c o r t e x o f t h e cotyledon,
the c o r t e x of the t r a n s i t i o n r e g io n
b e t w e e n r o o t an d s h o o t ,
and o c c a s i o n a l l y
in the
139 ~i
e o to d e rm is of th e c o ty le d o n . 11.
P o l y p l o i d c e l l s w e re f o u n d t o c o n s t i t u t e a p a r t o f n o r m a l d e v e l o p m e n t of s e e d l i n g s o f A l l i u m s t e l l a t u m r a n g i n g i n l e n g t h from 8 mm. t o 25 mm.
12.
T e t r a p l o i d d i v i s i o n s were i n c l u d e d i n t h e n o r m a l developm ent o f s e e d l i n g s o f A llium f i s t u l o s u m t h a t w ere 10 t o 40 mm. l o n g .
13.
A llium schoenoprasum in c lu d e d p o l y p lo i d c e l l s i n th e n o r m a l d e v e l o p m e n t o f y ou ng s e e d l i n g s 10 t o 33 mm. long.
14.
P o l y p l o i d c e l l s w e r e f o u n d t o be i n c l u d e d i n t h e norm al developm ent of s e e d l i n g s of Allium validum t h a t r a n g e d b e t w e e n 10 and 45 mm. i n l e n g t h .
15.
When s e e d l i n g s o f A l l i u m p orru m w e re 10 t o 44 mm. lon g ,
t e t r a p l o i d c e l l s were in c l u d e d as a f a c t o r
in t h e i r normal d e v elo p m e n t, 16.
S e e d l i n g s o f A l l i u m ammophilum t h a t w e re 10 t o 43 mm. lo n g in c lu d e d p o l y p l o i d d i v i s i o n s i n t h e i r normal developm ent.
17.
P o l y p l o i d c e l l s w e re f o u n d t o c o n s t i t u t e a f a c t o r i n th e normal developm ent o f s e e d l i n g s of Allium t u b e r o s u m 10 t o 40 mm. l o n g .
18.
A b e r r a t i o n s such as b i n u c l e a t e c e l l s , shaped n u c l e i ,
dum b-bell
t e l o p h a s e b r i d g e s a nd chromosomal
f r a g m e n t s w ere f o u n d t o be p r e s e n t i n a l l used i n t h i s i n v e s t i g a t i o n .
species
140 19.
O ccasionally,
two m i c r o n u c l e i were f o u n d i n t h e same
c e l l as the m acronucleus. 2 0.
Thes e a b e r r a t i o n s were f o u n d t o e x i s t i n t h e same r e g i o n s as th e t e t r a p l o i d c e l l s .
2 1.
T h e r e i s p r o b a b l y no c o n n e c t i o n b e t w e e n t h e a b n o r m a l i t i e s and t h e p o l y p l o i d c e l l s .
22.
The f o r m a t i o n o f d u m b - b e l l s h a p e d n u c l e i and b i n u c l e a t e c e l l s i s a s s o c i a t e d w i t h chro m o som a l b r i d g e s and f r a g m e n t s ,
2 5.
There i s a r e l a t i o n betw een th e d i s a p p e a r a n c e of the p o l y p l o i d c e l l s and t h e f o r m a t i o n o f t h e f i r s t l e a f .
24.
The p o l y p l o i d c e l l s d i s a p p e a r b e f o r e t h e f i r s t l e a f appears m acro sco p ically .
25.
The c e l l s o f A l l i u m ammophllum a r e i n g e n e r a l l a r g e r th an th o se of the o t h e r s p e c ie s used in t h i s in v estig atio n .
26.
The t e t r a p l o i d c e l l s were l a r g e r t h a n t h e d i p l o i d c e l l s o f t h e same s p e c i e s .
2 7.
P o ly p lo id c e l l s o ccu rred w ith g r e a t e r frequency in some s p e c i e s .
2 8.
The f r e q u e n c y o f t h e a b e r r a t i o n s v a r i e d w i t h t h e sp ecies.
~i
141
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y ¥ itk u s , E .R .,
1945•
Spinacia.
E n d o m ito sis i n ta p e t& l c e l l s of
American J o u r n a l of B o ta n y .
_____________ , and C.A. B e r g e r ,
1947.
32:
326-330.
Polyploid m itosis
i n t h e n o r m a l d e v e l o p m e n t o f Mimosa p u d i c a . B u l l e t i n of the Torrey B o ta n ic a l C lub. W i t t e , M.Q ., 1 9 4 7 .
74:
279-282.
C o m p a r a t i v e c y t o l o g i c a l s t u d y of
th r e e s p e c ie s of C henopodiacea. T orrey B o t a n i c a l Club.
74:
B u l l e t i n of the
443-452.
VITA
J a n e K i n s l o w Me E l l i g o t t , d a u g h t e r of S a r a h C e c e l i a and Thomas F r a n c i s Me E l l i g o t t was b o r n i n New Ha ven, C o n n e c t i c u t ,
on
December 8 , 1 9 2 2 . She a t t e n d e d W a t e r b u r y C a t h o l i c High S c h o o l from w h i c h sh e was g r a d u a t e d w i t h h o n o r in J u n e , 1939. I n S e p t e m b e r , 1 9 3 9 , she e n t e r e d Marymount C ollege.
She r e c e i v e d t h e d e g r e e o f B a c h e l o r
of A rts i n J u n e , 1943. I n S e p t e m b e r , 1945, s h e was a c c e p t e d i n t h e G r a d u a t e S c h o o l o f A r t s and S c i e n c e s o f Fordham U n i v e r s i t y f r o m w h i c h she r e c e i v e d t h e d e g re e of M aster of S cience in J u n e , 1946. She was a c c e p t e d a s a c a n d i d a t e f o r t h e P h .D . degree in J u l y ,
1946.