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Epidermal Studies in Amaranthus caudatus L., A. hypochondriacus L. and their F1 Hybrid
Flora, Bd. 167, S. 141-146 (1978)
Epidermal Studies in Amaranthu8 caudatu8 L., A. hypochondriacu8 L. and their Fl Hybrid M.
YUNUS,
K.
RASTOGI
(Mrs.), K. J.
AHMAD
and M.
PAL
National Botanic Gardens, Lucknow, India
Summary Foliar epidermal characters of Amaranthua caudatu8, A. hypochondriacua and their Fl hybrid have been studied. While the two parent species and their Fl hybrid show broad similarity in their epidermal features, the latter reveals closer affinity with A. hypochondriacua (paternal parent). This supports the earher findings which indicate the dominance of A. hypochondriacua on the basis of morphological studies.
Introduction The genus Amaranthus is widely distributed throughout tropical and temperate regions. It contains species that are cultivated for grains, as pot herbs and as ornamentals (SAUER 1967, PAL and KHOSHOO 1973). There is good deal of disagreement regarding the number of species within the genus which rangeCJ from 45-100 (SCHINZ 1934, AELLEN 1961). Much of the disagreement stems from the difficult synonymy which in turn has been complicated by frequent hybridization. The undue emphasis given by the earlier taxonomists to the variable characters like plant height, shape of the inflorescence and diversity in colour has added to the existing confusion. Foliar epidermal characterr, have often proved to be of taxonomic significance and have thus helped in proper taxonomic and phylogenetic evaluation of closely related taxa (see review STACE 1965, SINCLAIR and SHARMA 1971). Detailed study of the epidermal features in two species of the genus (A. caudatu8 and A. hypoclwndriacus) and one experimentally raised F 1 hybrid were undertaken to gain a better understanding of the interspecific relationsphips in the genus.
Materials and Methods Amctranthua caudatu8 ssp. caudatu8 and A. hypochondriacua both belong to the grain group species of the genus. In hybrid (A. caudatuaxA. hypochondriacua), A. caudatua ssp. caudatua and A. hypochondriacua were used as the maternal and paternal parents respectively. Material used in the present study was obtained from the plants raised from seeds. Methodology used in the preparation of foliar epidermal peels is after that of AHMAD (1974).
Observations Lower epidermis Intercor,tal cells irregular in shape, thin and sinuous walled lFigs, 1, 3, 5). Costal cells elongate, walls straight or arcuate, arranged in rows (Fig. 7), cells of minor veins slightly sinuous in A. caudatus and A. hypoclwndriacus (Fig. 8). Marginal cells elongate, thin and straight walled, arranged in rows (Fi{;. 10).
_Z2iJ t J -.}. LiE
142
Jj
:M. YVXlCS, K. RASTOGI, K. J.
AHMAD
and :U.
PAL
Stomata confined to intercostal areas (rarely on minor veins in A. caudatus (Fig. 9), anomo- and anisocytic in the same epidermal surface, rarely diacytic stomata present in A. caudahts and A. hypoLhondriacus; paracytic stomata rarely recorded in A. caudatus X A. hypochGndriaLus. Contiguous stomata sparse, arranged laterally (juxtaposed) or rarely associated at their poles (superposed), or with intermediate stages (Figs. 11-13); stomata with single guard cell (Fig. 14) or stomata with both the guard cells aborted (Fig. 15) rare in A. caudatus and A. hypochondriacus and spar'>e in A. caudatus X A. hypochondriacus. Polar thickenings of varying shape common (Figs. 18-27). Non-glandular hairs sparse in costal, intercostal and marginal areas (more common in costal areas of A. caudatus); uniseriate, rarely basal part biseriate; of two distinct types: (i) Hairs with club-shaped apex (Fig. 29), generally arising from a single epidermal cells (hair base 1-2 celled in A. caudatus, and A. hypochondriacus~and ... 1-3
E
r IH 17, ---{
vt--
100}lM
Figs. 1-6. Intercostal epidermal cells, 1. A. caudatu8, lower; 2. same upper; 3. A. hypochondriacua, lower; 4. same upper; 5. A. caudatuaxA.hypochondriucua, lower; 6. same upper; 7. costal cells in A. hypochondriucua; 8. cells of minor costae in A. caudatua; 9. stoma on minor vein of a costae in A. caudatua; 10. marginal cells of A. caudatua x A. hypochondriucua.
"'--"""
Epidermal Studies in Arna1'((nthus caudatu8 L.
"-,
.... ....... ".~
~-------.-...-
143
in A. caudatu8 X A. hypo~hondriacu8); co.mmo.nly 3-celled in length (1-6 celled in A. caudatu8 and A. hypochondriacus and 1-3 in A. caudatu8 X A. hypochondriac,u8). (ii) Hairs with narro.w tapering (Fig. 32) o.r blunt (Fig. 31) apex, co.mmo.nly arising fro.m a sin6le epidermal cell (hair base 1-8 celled in A. caudatus, 1-2 in A. hypochondriacus and 1-4 in A. caudatus X A. "h-ypochondriacus). Co.mmo.nly 4·celled in length (3-9 in A. wudatu8, 2-5 in A. hypochondriacu8 and 1-6 in A. caudatu8 X A. hypochondriacus). Upper epidermis Interco.stal cells po.lygo.nal in shape, thin, and straight o.r arcuate walled (Figs. 2, 4 & 6). Co.stal and marginal cells similar to. tho.se fo.und in the lo.wer epidermis. Sto.matd. ano.mo.- and aniso.cytic, rarely paracytic ill A. caudatu8 X A. hypochondriaCU8. Distributio.n, frequency o.f abno.rmal stomata (co.ntiguo.us, single guard cell o.r bo.th the guard cells abo.rted), and polar thickenings same as in lo.wer epidermis.
Figs. II-13. Contiguous stomata. 11. A. caudatusxA. hypoclwndriacus; 12. A. caudatus; 13. A. hypoclwndriacus}; 14. Single guard cell of A. caudatus; 15. Stoma with both the guard cells aborted in A. caudatus; 16 & 17. Stomata showing cytoplasmic contents in A. caudatus; IS-27. Stomata showing various types of polar thickenings (IS, 22, 23 & 25 in A. caudatus; 20, 21, 26 & 27 in A. hypochondriacus; 19 & 24 in A. caudatusXA. hypoclwndriacus); 2S-32. Non-glandular hairs (2S, 30 & 31. A. caudatusxA. hypochondriacus; 29 & 32. A. caudatus).
.. -
; ¢
144
M. YUl'ms, K.
RASTOGI.
K. J.
AHMAD
and M.
PAL
Non-glandular hairs sparse in costal, intercostal and marginal areas, uniseriate, rarely basal part biseriate; of two distinct types: (i) Hairs with club-shaped apex (Fig. 30), generally arising from a single epidermal cell (hair base 1-3 celled in A_ caudatus and A. caudatus X A_ hypor,hondriacus),; commonly 3-celled in length (1-5 celled in A. caudatuo and A. caudatus X A. hYPoCJhondriacus and 1-3 in A. hypochondriacus). (ii) Hairs with narrow tapering (Fig. 28) or blunt apex, arising from a single epidermal cell (hair base 1-4 celled in A. caudatus, 1-3 in A. hyp()chondriacu8 and 1-9 in A. caudatus X A. hypochondriacus); commonly 3-celled in length (1-4 in A. caudatus, 1-5 in A. hypochondriacus and A. caudatu8 X A. hydochondriacus).
Discussion The leaves in the hybrid are distorted and appear as if affected by mosaic virus. Associated with this are developmental abnormalities in other parts, which have been described in detail by PAL and KHOSHOO (1972). The,;e developmental defects are due to the geaic dishormony between the parental genomes and thus are genetic in nature. In their epidermal characters, the two parent species of Amaranthu8 (A. caudatu8 and A. hypochondriaous) and their F 1 hybrid (A. caudatu8 X A. hypochondriacus) Table 1. Measurements of epidermal characters Chcracters studied
A. hypochondriacu8
A. caudatus x A. hypochondriacu8
Upper
Lower
Upper
Lower
Upper
86
69
62
63
54
31.3% 67.5% 1.2%
39.7% 60.3%
43.9% 54.9% 1.2%
62.8% 37.2%
38.0% 60.8%
37.8% 61.0%
Size of stomata - pm
25x 21
23 x 19
34x 26
27 X 20
32x 25
30x
% of abnormal stomata
1.2
3.7
1.1
1.7
4.6
6.7
Stomatal index
24.0
14.8
26.3
20.8
18.8
12.5
Size of epidermal cell - pm
85x32
82x41
145 X 60
120x 49
141 X 44
103 X 43
74-96-148
48-66·108
82-113-168
74-89-131
115-170-217
102-136-176
A. caudatu8 Lower
Frequency of 153 stomata/mm sq. Type of stomata: Aniso Anomo Dia Para
Size of hair -
1.2%
1.2% ~5
{tm
Type I
82-120-143
82-134-180
Type II
82-205-668
120-134-189 102-160-234 60-92-116
...... -""."
Epidermal Studies in Amaranthu8 caudatu8 L.
..
~''"------.
145
show broadly similar features like irregular, sinuous walled intercostal cells of the lower epidermis and polygonal straight walled cells of the upper. Costal and marginal cells are also similar. Stomata are confined to intercostal areas and both anomocytic and anisocytic types ue present. Though the anomocytic "tomata are more common than the anisocytic ones, the percentage of anisocytic stomata is conspicuously higher in upper epidermis of A. hypochondriacUv (Table 1). Stomatal abilormalities like single guard cells, aborted guard cells, and contiguous stomata are rarely recorded. Non-glan~ular hairs of two distinct types are present and are quite similar in the three investigated taxa. There are, however, conspicuous differences, both qualitative and quantitative, which point towards closer affinities between one or the other parent on one hand, and the hybrid on the other. There are still some features, though minor ones, in which hybrid appears to be quite distinct from either of the two parents. In general Fl hybrid shows more similarity with A. hypochondriacus in its epidermal characters. These include: the size of epidermal cells and size and frequency of stomata (Table 1). In both of them, costal cells are straight-walled and stomata are confined to intercostal areas while in A. caudatus, cells of the minor veins are slightly sinuous walled and stomata are rarely present on minor veins also. Fl hybrid and A. wud6.tus, however, show considerable similarity in their upper epidermal hairs ttype ii with club-shaped apex) as they are 1-5 celled in length with 1-2 celled hair-base while in A. hypochondriacus they always arise from a single epidermal cell and are 1-3 celled in length. In several features Fl hybrid (A. caudatus X A. hypochondriacus) appears to be quite distinct from either of the two parent species. While diacytic stomata are recorded in A. caudatus and A. hypochondriacus, they are absent iil the hybrid; the paracytic type is present only in the hybrid. The percentage of abnormal stomata is also consid.erably higher in the hybrid than in the two parent species. The overall similarity in the epidermal features of the Fl hybrid with those of A. hypochondriacus (paternal parent) appears to be an expression of dominance of the latter vis-a-vis A. caudatus ssp. caudatus. Such an expression of dominance of A. hypochondriacus in respect of other morphological features as well has been reported by Pal and Khoshoo (1973) in the said hybrid. The precise reasons for the appearance of entirely new features like the paracytic stomata in the hybrid, not observed in either parent are not known at present, unless these are an indication of abnormal development itself.
Acknowledgements Our grateful thanks are due to Dr. T. N. KHOSHOO, Director, National Botanic Gardens for taking keen interest in the problem and also for providing necessary facilities. We are also thankful to our colleagues Mr. D. B. SHUKLA, Mr. BANSHI LAL and Mrs. K. SAXENA for their help in the collection of material and preparation of slides.
"'~i".,
146
•• •__ ••• i"
,. ,
_
:'I\. Yr:-;1::s, K. RASTOGI, K. J. AH3IAD and:'lI. PAL, Epidermal Studies in AmaI'Cmthu8
References AELLEN, P.: AmaranthacBae. In: G. HEGI, Illustrierte Flora von Mitteleuropa. 2. Auf!., Bd. Ill/2, S. 461-532. Munchen 1967. AHMAD, K. J.: Cuticular studies in some Nel8onioideae (Acanthaceae). Bot. J. Linn. Soc. 68, 73-80 (1974). PAL, M., and KHOSHOO, T. N.: Evolution and improvement of cultivated amaranths. V. Inviability, weakness and sterility in hybrids. J. Heredity 63, 78-82 (1972). _ _ Evolution and improvement of cultivated amaranths. VI. Cytogenetic relationship in grain types. Theoret. Applied Genetics 43, 242-251 (1973). SAUER, J. D.: The grain amaranths and their relatives: A revised taxonomic and geographic survey. Ann. Missouri. Bot. Gard. 54, 103-137 (1967). SCIDNZ, H.: Amaranthaceae. In: "Die Natiirlichen Pflanzenfamilien", (eds.: A. ENGLER and K. PRANTL, 16 C, 7-85 (1934). SINCLAIR, C. B., and SHARMA, G. K.: Epidermal and cuticular studies of plants. J. Tennessee Acad. Sci. 46, 2-11 (1971). STACE, C. A.: Cuticular studies as an aid to plant anatomy. Bull. B. M. (N. H.) Bot. 4, 1-78 (1965). Received August 31, 1977. Authors' address: Dr. M. YUNUS, (Mrs.), K. RASTOGI and Dr. K. J . .AHlIIAD, Plant Anatomy Laboratory, National Botanic Gardens, Lucknow 226 001, India, and Dr. M. PAL, Cytogenetics Laboratory, National Botanic Gardens, Lucknow 226001, India.
Epidermal Studies in Amaranthu8 caudatu8 L., A. hypochondriacu8 L. and their Fl Hybrid M.
YUNUS,
K.
RASTOGI
(Mrs.), K. J.
AHMAD
and M.
PAL
National Botanic Gardens, Lucknow, India
Summary Foliar epidermal characters of Amaranthua caudatu8, A. hypochondriacua and their Fl hybrid have been studied. While the two parent species and their Fl hybrid show broad similarity in their epidermal features, the latter reveals closer affinity with A. hypochondriacua (paternal parent). This supports the earher findings which indicate the dominance of A. hypochondriacua on the basis of morphological studies.
Introduction The genus Amaranthus is widely distributed throughout tropical and temperate regions. It contains species that are cultivated for grains, as pot herbs and as ornamentals (SAUER 1967, PAL and KHOSHOO 1973). There is good deal of disagreement regarding the number of species within the genus which rangeCJ from 45-100 (SCHINZ 1934, AELLEN 1961). Much of the disagreement stems from the difficult synonymy which in turn has been complicated by frequent hybridization. The undue emphasis given by the earlier taxonomists to the variable characters like plant height, shape of the inflorescence and diversity in colour has added to the existing confusion. Foliar epidermal characterr, have often proved to be of taxonomic significance and have thus helped in proper taxonomic and phylogenetic evaluation of closely related taxa (see review STACE 1965, SINCLAIR and SHARMA 1971). Detailed study of the epidermal features in two species of the genus (A. caudatu8 and A. hypoclwndriacus) and one experimentally raised F 1 hybrid were undertaken to gain a better understanding of the interspecific relationsphips in the genus.
Materials and Methods Amctranthua caudatu8 ssp. caudatu8 and A. hypochondriacua both belong to the grain group species of the genus. In hybrid (A. caudatuaxA. hypochondriacua), A. caudatua ssp. caudatua and A. hypochondriacua were used as the maternal and paternal parents respectively. Material used in the present study was obtained from the plants raised from seeds. Methodology used in the preparation of foliar epidermal peels is after that of AHMAD (1974).
Observations Lower epidermis Intercor,tal cells irregular in shape, thin and sinuous walled lFigs, 1, 3, 5). Costal cells elongate, walls straight or arcuate, arranged in rows (Fig. 7), cells of minor veins slightly sinuous in A. caudatus and A. hypoclwndriacus (Fig. 8). Marginal cells elongate, thin and straight walled, arranged in rows (Fi{;. 10).
_Z2iJ t J -.}. LiE
142
Jj
:M. YVXlCS, K. RASTOGI, K. J.
AHMAD
and :U.
PAL
Stomata confined to intercostal areas (rarely on minor veins in A. caudatus (Fig. 9), anomo- and anisocytic in the same epidermal surface, rarely diacytic stomata present in A. caudahts and A. hypoLhondriacus; paracytic stomata rarely recorded in A. caudatus X A. hypochGndriaLus. Contiguous stomata sparse, arranged laterally (juxtaposed) or rarely associated at their poles (superposed), or with intermediate stages (Figs. 11-13); stomata with single guard cell (Fig. 14) or stomata with both the guard cells aborted (Fig. 15) rare in A. caudatus and A. hypochondriacus and spar'>e in A. caudatus X A. hypochondriacus. Polar thickenings of varying shape common (Figs. 18-27). Non-glandular hairs sparse in costal, intercostal and marginal areas (more common in costal areas of A. caudatus); uniseriate, rarely basal part biseriate; of two distinct types: (i) Hairs with club-shaped apex (Fig. 29), generally arising from a single epidermal cells (hair base 1-2 celled in A. caudatus, and A. hypochondriacus~and ... 1-3
E
r IH 17, ---{
vt--
100}lM
Figs. 1-6. Intercostal epidermal cells, 1. A. caudatu8, lower; 2. same upper; 3. A. hypochondriacua, lower; 4. same upper; 5. A. caudatuaxA.hypochondriucua, lower; 6. same upper; 7. costal cells in A. hypochondriucua; 8. cells of minor costae in A. caudatua; 9. stoma on minor vein of a costae in A. caudatua; 10. marginal cells of A. caudatua x A. hypochondriucua.
"'--"""
Epidermal Studies in Arna1'((nthus caudatu8 L.
"-,
.... ....... ".~
~-------.-...-
143
in A. caudatu8 X A. hypo~hondriacu8); co.mmo.nly 3-celled in length (1-6 celled in A. caudatu8 and A. hypochondriacus and 1-3 in A. caudatu8 X A. hypochondriac,u8). (ii) Hairs with narro.w tapering (Fig. 32) o.r blunt (Fig. 31) apex, co.mmo.nly arising fro.m a sin6le epidermal cell (hair base 1-8 celled in A. caudatus, 1-2 in A. hypochondriacus and 1-4 in A. caudatus X A. "h-ypochondriacus). Co.mmo.nly 4·celled in length (3-9 in A. wudatu8, 2-5 in A. hypochondriacu8 and 1-6 in A. caudatu8 X A. hypochondriacus). Upper epidermis Interco.stal cells po.lygo.nal in shape, thin, and straight o.r arcuate walled (Figs. 2, 4 & 6). Co.stal and marginal cells similar to. tho.se fo.und in the lo.wer epidermis. Sto.matd. ano.mo.- and aniso.cytic, rarely paracytic ill A. caudatu8 X A. hypochondriaCU8. Distributio.n, frequency o.f abno.rmal stomata (co.ntiguo.us, single guard cell o.r bo.th the guard cells abo.rted), and polar thickenings same as in lo.wer epidermis.
Figs. II-13. Contiguous stomata. 11. A. caudatusxA. hypoclwndriacus; 12. A. caudatus; 13. A. hypoclwndriacus}; 14. Single guard cell of A. caudatus; 15. Stoma with both the guard cells aborted in A. caudatus; 16 & 17. Stomata showing cytoplasmic contents in A. caudatus; IS-27. Stomata showing various types of polar thickenings (IS, 22, 23 & 25 in A. caudatus; 20, 21, 26 & 27 in A. hypochondriacus; 19 & 24 in A. caudatusXA. hypoclwndriacus); 2S-32. Non-glandular hairs (2S, 30 & 31. A. caudatusxA. hypochondriacus; 29 & 32. A. caudatus).
.. -
; ¢
144
M. YUl'ms, K.
RASTOGI.
K. J.
AHMAD
and M.
PAL
Non-glandular hairs sparse in costal, intercostal and marginal areas, uniseriate, rarely basal part biseriate; of two distinct types: (i) Hairs with club-shaped apex (Fig. 30), generally arising from a single epidermal cell (hair base 1-3 celled in A_ caudatus and A. caudatus X A_ hypor,hondriacus),; commonly 3-celled in length (1-5 celled in A. caudatuo and A. caudatus X A. hYPoCJhondriacus and 1-3 in A. hypochondriacus). (ii) Hairs with narrow tapering (Fig. 28) or blunt apex, arising from a single epidermal cell (hair base 1-4 celled in A. caudatus, 1-3 in A. hyp()chondriacu8 and 1-9 in A. caudatus X A. hypochondriacus); commonly 3-celled in length (1-4 in A. caudatus, 1-5 in A. hypochondriacus and A. caudatu8 X A. hydochondriacus).
Discussion The leaves in the hybrid are distorted and appear as if affected by mosaic virus. Associated with this are developmental abnormalities in other parts, which have been described in detail by PAL and KHOSHOO (1972). The,;e developmental defects are due to the geaic dishormony between the parental genomes and thus are genetic in nature. In their epidermal characters, the two parent species of Amaranthu8 (A. caudatu8 and A. hypochondriaous) and their F 1 hybrid (A. caudatu8 X A. hypochondriacus) Table 1. Measurements of epidermal characters Chcracters studied
A. hypochondriacu8
A. caudatus x A. hypochondriacu8
Upper
Lower
Upper
Lower
Upper
86
69
62
63
54
31.3% 67.5% 1.2%
39.7% 60.3%
43.9% 54.9% 1.2%
62.8% 37.2%
38.0% 60.8%
37.8% 61.0%
Size of stomata - pm
25x 21
23 x 19
34x 26
27 X 20
32x 25
30x
% of abnormal stomata
1.2
3.7
1.1
1.7
4.6
6.7
Stomatal index
24.0
14.8
26.3
20.8
18.8
12.5
Size of epidermal cell - pm
85x32
82x41
145 X 60
120x 49
141 X 44
103 X 43
74-96-148
48-66·108
82-113-168
74-89-131
115-170-217
102-136-176
A. caudatu8 Lower
Frequency of 153 stomata/mm sq. Type of stomata: Aniso Anomo Dia Para
Size of hair -
1.2%
1.2% ~5
{tm
Type I
82-120-143
82-134-180
Type II
82-205-668
120-134-189 102-160-234 60-92-116
...... -""."
Epidermal Studies in Amaranthu8 caudatu8 L.
..
~''"------.
145
show broadly similar features like irregular, sinuous walled intercostal cells of the lower epidermis and polygonal straight walled cells of the upper. Costal and marginal cells are also similar. Stomata are confined to intercostal areas and both anomocytic and anisocytic types ue present. Though the anomocytic "tomata are more common than the anisocytic ones, the percentage of anisocytic stomata is conspicuously higher in upper epidermis of A. hypochondriacUv (Table 1). Stomatal abilormalities like single guard cells, aborted guard cells, and contiguous stomata are rarely recorded. Non-glan~ular hairs of two distinct types are present and are quite similar in the three investigated taxa. There are, however, conspicuous differences, both qualitative and quantitative, which point towards closer affinities between one or the other parent on one hand, and the hybrid on the other. There are still some features, though minor ones, in which hybrid appears to be quite distinct from either of the two parents. In general Fl hybrid shows more similarity with A. hypochondriacus in its epidermal characters. These include: the size of epidermal cells and size and frequency of stomata (Table 1). In both of them, costal cells are straight-walled and stomata are confined to intercostal areas while in A. caudatus, cells of the minor veins are slightly sinuous walled and stomata are rarely present on minor veins also. Fl hybrid and A. wud6.tus, however, show considerable similarity in their upper epidermal hairs ttype ii with club-shaped apex) as they are 1-5 celled in length with 1-2 celled hair-base while in A. hypochondriacus they always arise from a single epidermal cell and are 1-3 celled in length. In several features Fl hybrid (A. caudatus X A. hypochondriacus) appears to be quite distinct from either of the two parent species. While diacytic stomata are recorded in A. caudatus and A. hypochondriacus, they are absent iil the hybrid; the paracytic type is present only in the hybrid. The percentage of abnormal stomata is also consid.erably higher in the hybrid than in the two parent species. The overall similarity in the epidermal features of the Fl hybrid with those of A. hypochondriacus (paternal parent) appears to be an expression of dominance of the latter vis-a-vis A. caudatus ssp. caudatus. Such an expression of dominance of A. hypochondriacus in respect of other morphological features as well has been reported by Pal and Khoshoo (1973) in the said hybrid. The precise reasons for the appearance of entirely new features like the paracytic stomata in the hybrid, not observed in either parent are not known at present, unless these are an indication of abnormal development itself.
Acknowledgements Our grateful thanks are due to Dr. T. N. KHOSHOO, Director, National Botanic Gardens for taking keen interest in the problem and also for providing necessary facilities. We are also thankful to our colleagues Mr. D. B. SHUKLA, Mr. BANSHI LAL and Mrs. K. SAXENA for their help in the collection of material and preparation of slides.
"'~i".,
146
•• •__ ••• i"
,. ,
_
:'I\. Yr:-;1::s, K. RASTOGI, K. J. AH3IAD and:'lI. PAL, Epidermal Studies in AmaI'Cmthu8
References AELLEN, P.: AmaranthacBae. In: G. HEGI, Illustrierte Flora von Mitteleuropa. 2. Auf!., Bd. Ill/2, S. 461-532. Munchen 1967. AHMAD, K. J.: Cuticular studies in some Nel8onioideae (Acanthaceae). Bot. J. Linn. Soc. 68, 73-80 (1974). PAL, M., and KHOSHOO, T. N.: Evolution and improvement of cultivated amaranths. V. Inviability, weakness and sterility in hybrids. J. Heredity 63, 78-82 (1972). _ _ Evolution and improvement of cultivated amaranths. VI. Cytogenetic relationship in grain types. Theoret. Applied Genetics 43, 242-251 (1973). SAUER, J. D.: The grain amaranths and their relatives: A revised taxonomic and geographic survey. Ann. Missouri. Bot. Gard. 54, 103-137 (1967). SCIDNZ, H.: Amaranthaceae. In: "Die Natiirlichen Pflanzenfamilien", (eds.: A. ENGLER and K. PRANTL, 16 C, 7-85 (1934). SINCLAIR, C. B., and SHARMA, G. K.: Epidermal and cuticular studies of plants. J. Tennessee Acad. Sci. 46, 2-11 (1971). STACE, C. A.: Cuticular studies as an aid to plant anatomy. Bull. B. M. (N. H.) Bot. 4, 1-78 (1965). Received August 31, 1977. Authors' address: Dr. M. YUNUS, (Mrs.), K. RASTOGI and Dr. K. J . .AHlIIAD, Plant Anatomy Laboratory, National Botanic Gardens, Lucknow 226 001, India, and Dr. M. PAL, Cytogenetics Laboratory, National Botanic Gardens, Lucknow 226001, India.