Meiotic behaviour and pollen fertility in the seventeen

Vol. 55, no. 4: 341-347, 2002
CARYOLOGIA
Meiotic behaviour and pollen fertility in the seventeen
Brazilian species of Adesmia DC. (Leguminosae)
SOLANGE BOSIO TEDESCO1, MARIA TERESA SCHIFINO-WITTMANN*, 2 and MIGUEL DALL’AGNOL2
1
Departamento de Biologia, Centro de Ciências Naturais e Exatas, Universidade Federal de Santa Maria, 97105-900 Santa
Maria, RS, Brazil.
2 Departamento de Plantas Forrageiras e Agrometeorologia, Faculdade de Agronomia, UFRGS. Caixa Postal 776. 91501-970
Porto Alegre, RS, Brazil.
Abstract - The southern region of Brazil, especially Rio Grande do Sul State, is
famous for its natural pastures. Many of the native legume species are potentially good forages. Among them are some of the 17 diploid (2n=20) species and one
variety of Adesmia DC described for Brazil. Meiotic behaviour, meiotic indexes
and pollen fertility were studied in 38 accessions of A. araujoi, A. arillata, A. bicolor, A. ciliata, A. incana (including one tetraploid accession), A. latifolia, A. muricata, A. psoraleoides, A. punctata, A. reitziana, A. riograndensis, A. rocinhensis,
A. securigerifolia, A. tristis and A. vallsii. For A. paranensis and A. sulina only
pollen data were available. Meiotic behaviour was essentially regular in most of
the accessions, with 10 bivalents (II) at diakinesis and metaphase I and regular
chromosome segregation at anaphase and telophase I and II. Uni, tri and quadrivalents were sometimes observed at diakinesis and metaphase I as well as laggards
and bridges at anaphase and telophase I and II. Meiotic indexes and pollen fertility were normally over 90% or nearly 90%. However, in some accessions up to
30% of the cells presented abnormalities in chromosome pairing, chromosome
segregation or low meiotic indexes or pollen fertility. From a taxonomic point of
view neither chromosome number nor meiotic behaviour can be used to distinguish between the taxa analysed. Regarding an applied approach, as for example
in planning crosses or in seed production for cultivated pastures, the absence of
major meiotic anomalies and the high pollen fertility is an advantage.
Key Words: Adesmia, forages, meiotic behaviour, pollen fertility, taxonomy.
INTRODUCTION
The southern region of Brazil, especially Rio
Grande do Sul State, is famous for its native pastures, formed mainly by warm season species, the
basis of cattle feeding. Due to a seasonal lack of
forage during the winter months there is a search
of native grass and legume species that could be
intensively used as forages, directly or after selection and breeding procedures. One of the most
promising genus is Adesmia DC. (Leguminosae,
Faboideae, Adesmieae) which comprises around
* Corresponding author: fax ++55 513 316 6045; e-mail:
[email protected].
200 annual and perennial, herbaceous and shrubby species endemic to South America, distributed along the Andes mountains, from northern
Peru to Tierra del Fuego. The genus center of origin is supposed to be central Chile and western
Argentina (BURKART 1967).
Seventeen species and one variety are found in
Brazil, all of them restricted to the southern part
of the country, including Rio Grande do Sul, Santa Catarina and Paraná States (MIOTTO and
LEITÃO-FILHO 1993). Several species of Adesmia
from Southern Brazil present good growth during
winter, are well adapted to the region environmental conditions, are widespread and have a
high nutritional value. For example, A. latifolia
342
(Spreng.) Vog. has a good forage yield potential
as well as high mineral content (SCHEFFER-BASSO
et al. 2000, 2001). Crude protein percentages
range from 6.9 to 17.5 in A. tristis Vog., 18.6 in
A. latifolia, 17.9 to 21.5 in A. ciliata Vog., 19.7 in
A. psoraleoides Vog. and 23.4 in A. punctata
(Poir.) DC. (DALL’AGNOL and GOMES 1994).
High crude protein, as well as in vitro organic
matter digestibility values were also found for A.
latifolia, A. tristis and A. punctata by SCHEFFERBASSO et al. (2000, 2001).
Regarding mode of reproduction, A. latifolia
is a versatile species, allowing self and cross fertilization (TEDESCO et al. 1998) as well as A. bicolor (Poir.) DC., A. muricata (Jacq.) DC., A. punctata and A. riograndensis Miotto, while A. incana
Vog. reproduces by self-pollination and A. tristis
mainly by cross-pollination. (TEDESCO et al.
2000a). There is strong evidence that A. securigerifolia Hert. is a self-pollinating species and A. arillata Miotto, A. ciliata, A. psoraleoides, A. rocinhesis Burk., A. reitziana Burk., A. sulina Miotto
and A. vallsii Miotto are probably cross-pollinated (TEDESCO et al. 2000a).
Cytogenetic studies in Adesmia are few. From
literature data, chromosome numbers are known
for less than 20% of the total number of species
(FEDOROV 1969; GOLDBLATT 1981, 1984, 1985,
1988; GOLDBLATT and JOHNSON 1990, 1991,
1994, 1996, 1998, 2000). Most of the studied
species are diploid with 2n=20 chromosomes,
and a few are tetraploid (2n=40). Since the first
chromosome number determinations in the
genus, n=10 has been suggested as the genus
basic number (CASTRONOVO 1945; COVAS and
SCHNACK 1946; COVAS 1949; KRAPOVICKAS and
KRAPOVICKAS 1951; COVAS and HUNZIKER 1954;
RAHN 1960; HUNZIKER et al. 1985).
For Brazilian species, chromosome numbers
have been determined (MORAES-FERNANDES and
BARRETO 1964; MIOTTO and FORNI-MARTINS
1995; COELHO 1996; COELHO and BATTISTIN
1998; TEDESCO et al. 2000b). Observations on
chromosome morphology for some species
(COELHO 1996), showed that most have small
(ca. 1.5 to 2.5µm) meta and sub-metacentric
chromosomes, with suggestions of inter-specific
differences on the total chromosome complement length. Previous observations on meiotic
behaviour of 12 accessions of seven species
showed regular chromosome pairing and high
meiotic indexes and pollen fertility (COELHO
1996; COELHO and BATTISTIN 1998).
TEDESCO, SCHIFINO-WITTMANN
and DALL’AGNOL
As can be seen from literature data, a detailed
and comparative analysis of meiotic behaviour
and pollen fertility in all the 17 Brazilian Adesmia
species was lacking, and therefore this was the
objective of the present paper.
This work is part of a multidisciplinary project
on germplasm characterisation of the Brazilian
Adesmia species, inserted in a broader research
line of characterisation, utilisation and preservation of native species of the natural pastures of
Rio Grande do Sul, developed at the Departamento de Plantas Forrageiras, Faculdade de
Agronomia, Universidade Federal do Rio Grande
do Sul, Brazil.
MATERIAL AND METHODS
Seeds from 41 accessions of the 17 Brazilian
species were collected in several places of Rio Grande
do Sul, Santa Catarina and Parana States (ca. 33º to
23º latitude South) (Table 1). Taxonomic vouchers of
the mother plants are kept at the ICN (Instituto de
Biociências, Universidade Federal do Rio Grande do
Sul, Brazil) and CENARGEM (Centro Nacional de
Recursos Genéticos e Biotecnologia - EMBRAPA,
Brazil) Herbaria.
The seeds were scarified with sand paper, germinated in petri dishes with moistened filter paper in a
growth chamber at 25º C and a 16h light- 8 h darkness
photoperiod at the Cytogenetics Laboratory of the
Universidade Federal de Santa Maria. The seedlings
were transferred to plastic cups with soil when they
reached 3 about cm height, and to 5 kg-capacity plastic pots when they were forty-five days old. The pots
were kept in a greenhouse at the Forestry Department, Universidade Federal de Santa Maria, where
the flower buds for meiotic analysis were collected.
Young flower buds were collected and fixed in 3:1
ethanol-acetic acid for 24 h at room temperature, and
afterwards transferred to 70% ethanol and kept at
4º C until slide preparation.
For meiosis analysis, the slides were prepared by
squashing the anthers in 1% acetic orcein. All observable phases of meiosis were analysed but a special
emphasis was given to chromosome associations at
diakinesis and metaphase I and chromosome segregation at anaphase and telophase I and anaphase and
telophase II. Meiotic indexes (mi) were calculated
according LOVE (1949), being considered as normal
those tetrads with four equal-sized cells, and as abnormal any other formations. Around 300 tetrads (200600) per accession were examined.
For pollen fertility (pf) estimation, the anthers were
stained with 2% propionic carmine. Ten slides from
different flowers generally from different inflores-
343
MEIOTIC BEHAVIOUR IN ADESMIA
Table 1 – List of the Adesmia species and accessions examined.
Species
A. araujoi
A. araujoi
A. arillata
A. arillata
A. bicolor
A. bicolor
A. bicolor
A. bicolor
A. ciliata
A. incana var. incana
A. incana
A. incana
A. incana
A. incana
A. latifolia
A. latifolia
A. muricata
A. muricata
A. muricata
A. paranensis
A. psoraleoides
A. psoraleoides
A. psoraleoides
A. psoraleoides
A. psoraleoides
A. punctata var. hilariana
A. punctata
A. reitziana
A. riograndensis
A. riograndensis
A. rocinhensis
A. rocinhensis
A. securigerifolia
A. securigerifolia
A. securigerifolia
A. securigerifolia
A. sulina
A. tristis
A. tristis
A. vallsii
A. vallsii
Accessiona
V 10730
SB s/n
V 11318
V 11346
V 9688
V 12243
V 12340
Mi 1512
V 8183
Te s/n
V 9636
V 9637
V 9654
V 10288
EEL 15025
Zm 1775
V 9570
Zm 236
V 10289
Te s/n
V 10760
V 11355
V 11425
Mi 1564
V 11325
V 6885
V 10812
Mi 1630
V 9590
Zm 419
V 7470
V 11507
V 6978
V 9615
Te s/n
Te s/n
V 11500
V 10766
V 10814
V 11439
V 11465
Codeb
BRA-000817
BRA-001473
BRA-000825
BRA-000841
BRA-000183
BRA-000892
BRA-000884
BRA-001520
BRA-000272
BRA-001503
BRA-000604
BRA-000311
BRA-000329
BRA-001015
BRA-001732
BRA-001414
BRA-000485
BRA-001040
BRA-001058
BRA-001716
BRA-001066
BRA-001082
BRA-001091
BRA-001724
BRA-001074
BRA-001104
BRA-001112
BRA-001708
BRA-000761
BRA-001139
BRA-000116
BRA-001163
BRA-000060
BRA-000621
BRA-001481
BRA-001490
BRA-001261
BRA-001317
BRA-001325
BRA-001368
BRA-001384
Place of collectionc
Soledade, RS
Passo Fundo, RS
Guarapuava, PR
Guarapuava, PR
Uruguaiana, RS
São Borja, RS
Dom Pedrito, RS
Bagé, RS
Lages, SC
Bagé, RS
Santana Livramento, RS
Santana Livramento, RS
Quaraí, RS
Bagé, RS
SC
Imbé, RS
Canguçú, RS
Caçapava do Sul, RS
Caçapava do Sul, RS
Santiago, RS
Lagoa Vermelha, RS
Guarapuava, PR
Abelardo Luz, SC
Vargem Bonita, SC
Guarapuava, PR
Vacaria, RS
Vacaria, RS
Urubici, SC
Santana da Boa Vista, RS
Caçapava do Sul, RS
São Joaquim, SC
Palmas, PR
Bagé, RS
Bagé, RS
Bagé, RS
Bagé, RS
Água Doce, SC
Vacaria, RS
Vacaria, RS
Palmas, PR
Palmas, PR
collector’s number
code number at EMBRAPA-CENARGEN
c PR-Paraná State; RS-Rio Grande do Sul State; SC-Santa Catarina State
a
b
cences were prepared per accession and the number of
stained (viable) and empty (sterile) grains were recorded (Fig. 1). At least 1000 pollen grains were examined
per plant.
RESULTS AND DISCUSSION
Meiotic behaviour, meiotic indexes and pollen
fertility were studied in 38 accessions of A. araujoi
Burk. A. arillata, A. bicolor, A. ciliata, A. incana,
A. latifolia, A. muricata, A. psoraleoides, A. punctata, A. reitziana, A. riograndensis, A. rocinhensis,
A. securigerifolia, A. tristis and A. vallsii. For
A. paranensis Burk., A. sulina and one accession of
A. psoraleoides only data on pollen fertility was
available (Table 2). All accessions examined were
diploid, 2n=20, except the tetraploid A. incana V
9637.
Meiotic behaviour was essentially regular in
most of the accessions, with 10 bivalents (II) at
diakinesis and metaphase I (Fig. 2) and normal
chromosome segregation at anaphase and
telophase I and II. Uni (I), tri (III) and quadrivalents (IV) were eventually observed as well as irregular segregation and laggards and bridges at
344
TEDESCO, SCHIFINO-WITTMANN
and DALL’AGNOL
Fig. 1 – Fertile (stained) and sterile (empty) pollen grains in
A. securigerifolia. Scale Bar 4µm.
Fig. 2 – Diakinesis with 10 II in A. arillata. Scale Bar 4 µm.
anaphase I and II. Meiotic indexes were normally
over 90%, indicating meiotically stable plants
(LOVE 1949), or nearly 90%. The abnormalities at
this stage included mostly dyads and triads. Pollen
fertility was also high, over 90%, for most accessions (Table 2).
The main deviations from normality were
observed in A. latifolia Zm 1775 (20% of cells
with trivalents and/or univalents), A. muricata V
9570 (15% of cells with trivalents and/or univalents), A. muricata Zm 236 (20% of the cells with
one quadrivalent), A. arillata V 11318 (mi 68.89,
pf 43.77), A. incana V 9636 (pf 76.76), A. muricata V 10289 (pf 61,13), A. paranensis Te s/n (pf
61.13), A. psoraleoides V 11325 (pf 13.47). The
only case in which a relation between meiotic
anormalities and rather low meiotic meiotic indexes could be suggested was A. ciliata V 8183 (30%
of the cells with univalents, 30% of cells with laggards at anaphase I or anaphase II, im 67,89)
although pollen fertility was over 90%. For A.
paranensis Te s/n and A. psoraleoides V 11325
only pollen fertility data were obtained. As meiotic
behaviour was not analysed in V 11325, it could
not be verified if the very low pollen fertility
(13.47) were due to irregularities during meiosis.
The tetraploid accession of A. incana (V 9637)
presented regular meiosis with 20II at metaphase
I, normal segregation at anaphase I and II and
very high meiotic index and pollen fertility
(Table 2), confirming a previous work by COELHO
and BATTISTIN (1998). Tetraploid accessions have
been reported in A. incana by other researchers
(CASTRONOVO 1945; HUNZIKER et al. 1985; MIOTTO and FORNI-MARTINS 1995). If this accession is
supposed to be of autotetraploid origin, as the
absence of any trait distinguishing it from the conspecific diploid accessions may suggest, strong
forces to a strict bivalent pairing, such as cytological diploidisation are expected to be acting, but
due to the lack of additional studies, any suggestions on this matter are purely speculative.
The results show that, apart from a few exceptions, most of the natural populations of the
Adesmia species studied have a regular meiotic
behaviour and high pollen fertility. From a taxonomic point of view neither chromosome number nor meiotic behaviour could be used to distinguish between the taxa analysed. Regarding an
applied approach, as for example in planning
crosses for breeding, or in seed production for
cultivated pastures, the absence of major meiotic
anomalies and the generally high pollen fertility is
an advantage.
Concluding remarks
Cytogenetic studies in Adesmia should be
intensified. The results reported in this paper are
345
MEIOTIC BEHAVIOUR IN ADESMIA
Table 2 - Meiotic associations, chromosome segregation, meiotic indexes and pollen fertility in Brazilian species of Adesmia.
Species
A. araujoi
A. araujoi
A. arillata
A. arillata
A. bicolor
A. bicolor
A. bicolor
A. bicolor
A. ciliata
A. incana
var. incana
A. incana
A. incana
A. incana
A. incana
A. latifolia
A. latifolia
A. muricata
A. muricata
A. muricata
A. paranensis
A. psoraleoides
A. psoraleoides
A. psoraleoides
A. psoraleoides
A. psoraleoides
A. punctata
var. hilariana
A. punctata
A. reitziana
A. riograndensis
A. riograndensis
A. rocinhensis
A. rocinhensis
A. securigerifolia
A. securigerifolia
A. securigerifolia
A. securigerifolia
A. sulina
A. tristis
A. tristis
A. vallsii
A. vallsii
Accessions
Diakinesis/Metaphase I
Anaphase I/TelophaseI
Anaphase II/Telophase II
number
of cells
chromosome
associations
number chromosome
of cells segregation
number
of cells
chromosome
segregation
33
16
48
10
60
59
63
41 (18)a
10 II
10 II
10 II
10 II
10 II
10 II
10 II
10 IIa
63
10
19
21
56
71
16
14
regular
regular
regular
regular
regular
regular
regular
regular
25
28
10
4
26
84
12
44 (13)b
regular
regular
regular
regular
regular
regular
regular
regular b
97.59
97.59
68.89
67.21
97.99
96.30
93.28
99.10
67.89
99.15
95.11
43.77
61.13
98.78
99.46
95.76
97.93
94.83
Te s/n
V 9636
V 9637
V 9654
V 10288
EEL 15025
Zm 1775
V 9570
Zm 236
V 10289
Te s/n
V 10760
V 11355
V 11425
Mi 1564
V 11325
19
159
62
48
11
98
53 (10)c
118 (20)d
29 (8)e
18
10 II
10 II
20 II
10 II
10 II
10 II
10 IIc
10 IId
10 IIe
10 II
30
156
44
12
23
36
32
17
6
14
regular
regular
regular
regular
regular
regular
regular
regular
regular
regular
12
31
25
30
42
11
24
54
15
18
regular
regular
regular
regular
regular
regular
regular
regular
regular
regular
94.93
91.50
96.67
88.43
95.94
95.65
91.15
80.69
75.72
98.36
40
10
7
115
10 II
10 II
10 II
10 II
14
18
regular
regular
28
8
regular
regular
23
regular
49
regular
86.30
98.00
100.00
93.55
99.22
76.76
99.64
99.61
97.56
98.95
97.89
92.98
90.11
97.19
67.49
98.01
97.99
97.22
92.61
13.47
V 6885
V 10812
Mi 1630
V 9590
Zm 419
V 7470
V 11507
V 6978
V 9615
Te s/n
(BRA 001481)
Te s/n
(BRA 001490)
V 11500
V 10766
V 10814
V 11439
V 11465
3
25
24
11
63
68
12
19
12
10 II
10 II
10 II
10 II
10 II
10 II
10 II
10 II
10 II
4
107(18)f
33
16
78
15
100
64
3
regular
regular
regular
regular
regular
regular
regular
regular
regular
33
55
10
regular
regular
regular
50
9
84
32
26
regular
regular
regular
regular
regular
82.41
81.08
100.00
94.79
84.50
97.98
91.92
97.16
19
10 II
88
regular
34
regular
93.26
10
10 II
6
regular
27
regular
95.48
25
32
20
36
10 II
10 II
10 II
10 II
24
30
12
26
regular
regular
regular
regular
19
33
13
30
regular
regular
regular
regular
96.93
96.72
98.25
99.14
V 10730
SB s/n
V 11318
V 11346
V 9688
V 12243
V 12340
Mi 1512
V 8183
18 cells with 1-4 univalents
13 cells with irregular segregation or 1-3 laggards
c 10 cells with one trivalent and/or 1-2 univalents
d 20 cells with one trivalent and/or 1-8 univalents
e 8 cells with univalents or quadrivalents
f 18 cells with irregular segregation or 1-2 laggards
a
b
Meiotic index Pollen fertility
(%)
(%)
99.76
86.10
82.60
99.24
99.68
90.74
87.58
98.69
98.30
96.78
99.63
98.89
97.57
97.32
346
TEDESCO, SCHIFINO-WITTMANN
practically non-existent for other species besides
the Brazilian ones. A broader survey of chromosome numbers, as well as the analysis of meiotic
behaviour and pollen fertility in other species
apart from the Brazilian taxa, and verification of
intraspecific variability in chromosome number
as such found in A. incana should be done to a
better cytogenetic characterization of the genus.
Especifically for the Brazilian species, an examination of more accessions, mainly of the poorly
analysed A. sulina and A. paranensis should be
undertaken.
Acknowledgments – To Dr. José Francisco
Montenegro Valls, EMBRAPA-CENARGEN for
most of the seedlots and for his continual support
for our work. To Dr. Alice Battistin and Dr. Juarez
Hope, Universidade Federal de Santa Maria, for allowing the utilisation of their laboratory and greenhouse facilities. To Moises Stefanello for his help in
the pollen analyis. To CNPq (Conselho Nacional de
Desenvolvimento Científico e Tecnológico), Brazil,
for finnancial support and to CAPES (Comissão de
Aperfeiçoamento de Pessoal de Ensino Superior),
Brazil for the PhD scholarship to the first author.
REFERENCES
BURKART A., 1967 – Sinópsis del género sudamericano de Leguminosas Adesmia DC. Darwiniana,
14: 463-568.
CASTRONOVO A., 1945 – Estudos cariologicos de
doce especies de leguminosas argentinas. Darwiniana, 7: 38-57.
COELHO L.G.M., 1996 – Citogenética e qualidade de
forragem de espécies de Adesmia DC. nativas no
Rio Grande do Sul. MSc Thesis, Universidade
Federal de Santa Maria, Brazil.
COELHO L.G.M. and BATTISTIN A., 1998 – Meiotic
behavior of Adesmia DC. (LeguminosaeFaboideae) species native to Rio Grande do Sul,
Brazil. Genetics and Molecular Biology, 21: 403406.
COVAS G., 1949 – Estudios cariológicos em antófitas,
III Parte. Darwiniana, 9: 158-162.
COVAS G. and HUNZIKER J.H., 1954 – Estudos cariológicos em antófitas. IV Parte. Revista de Investigaciones Agricolas, 8: 249-253.
COVAS G. and SCHNACK B., 1946 – Número de cromosomas em antófitas de la region Cuyo (República Argentina). Revista Argentina de Agronomia,
13: 153-166.
DALL’AGNOL M. and GOMES K.E., 1994 – Qualidade de acessos de leguminosas nativas do gênero
and DALL’AGNOL
Adesmia. Anais Reunião da Sociedade Brasileira
de Zooetcnia, 31: 653.
FEDOROV AN.A., 1969 – Chromosome Numbers of
Flowering Plants. Russian Academy of Sciences.
GOLDBLATT P., 1981 – Index to Plant Chromosome
Numbers 1975-1978. Monographs in Systematic
Botany, 5. Missouri Botanical Garden.
–, 1984 – Index to Plant Chromosome Numbers
1979-1981. Monographs in Systematic Botany, 8.
Missouri Botanical Garden.
–, 1985 – Index to Plant Chromosome Numbers
1982-1983. Monographs in Systematic Botany,
13. Missouri Botanical Garden.
–, 1988 – Index to Plant Chromosome Numbers
1984-1985. Monographs in Systematic Botany,
23. Missouri Botanical Garden.
GOLDBLATT P. and JOHNSON D.E., 1990 – Index to
Plant Chromosome Numbers 1986-1987. Monographs in Systematic Botany, 30. Missouri Botanical Garden.
–, 1991 – Index to Plant Chromosome Numbers
1988-1989. Monographs in Systematic Botany,
40. Missouri Botanical Garden.
–, 1994 – Index to Plant Chromosome Numbers
1990-1991. Monographs in Systematic Botany,
51. Missouri Botanical Garden.
–, 1996 – Index to Plant Chromosome Numbers
1992-1993. Monographs in Systematic Botany,
58. Missouri Botanical Garden.
–, 1998 – Index to Plant Chromosome Numbers
1994-1995. Monographs in Systematic Botany,
69. Missouri Botanical Garden.
–, 2000 – Index to Plant Chromosome Numbers
1996-1997. Monographs in Systematic Botany,
81. Missouri Botanical Garden.
HUNZIKER J.H., XIFERDA C.C. and WULFF A.F.,
1985 – Estudios cromosomicos en angiospermas
de sudamerica. Darwiniana, 26: 7-14.
KRAPOVICKAS A. and KRAPOVICKAS A.M., 1951 –
Notas citológicas sobre leguminosas. Darwiniana,
9: 612-613.
LOVE R.M., 1949 – La citologia como ayuda practica
al mejoramiento de los cereales. Revista Argentina Agronomia, 16: 1-13.
MIOTTO S.T.S. and FORNI-MARTINS E.R., 1995 –
Número cromossômico em espécies brasileiras de
Adesmia DC. (Leguminosae, Faboideae). Acta
botanica Brasilica, 8: 3-9.
MIOTTO S.T.S. and LEITÃO FILHO H.F., 1993 –
Leguminosae-Faboideae, Gênero Adesmia DC.
Boletim do Instituto de Biociências, 52: 1-157.
MORAES-FERNANDES M.I.B and BARRETO I., 1964 –
Estudos citotaxonômicos das pastagens nativas
do RGS. Anais Congresso Sociedade Botânica
do Brasil, 15.
MEIOTIC BEHAVIOUR IN ADESMIA
RAHN K., 1960 – Chromosome numbers in some
South American angiosperms. Botanisk
Tidsskrift, 56: 117-127.
SCHEFFER-BASSO S.M.S., JACQUES A.V.A., DALL’AGNOL M., RIBOLDI J. and CASTRO S.M.J.,
2000 – Dinâmica da formação de gemas, folhas e
hastes de espécies de Adesmia DC. e Lotus L.
Revista da Sociedade Brasileira de Zootecnia,
29: 1961-1968.
–, 2001 – Disponibilidade e valor nutritivo de forragem de leguminosas nativas (Adesmia DC.) e
exóticas (Lotus L.). Revista da Sociedade
Brasileira de Zootecnia, 30: 975-982.
TEDESCO S.B., DALL’AGNOL M. and SCHIFINOWittmann M.T., 1998 – Observações sobre o
347
modo de reprodução em Adesmia latifolia
Spreng.Vog. Ciência Rural, 28: 141-142.
T EDESCO S.B., D ALL ’A GNOL M., S CHIFINO WITTMANN M.T. and VALLS J.F.M., 2000a –
Mode of reproduction of Brazilian species of
Adesmia DC. (Leguminosae). Genetics and Molecular Biology, 23: 475-478.
TEDESCO S.B., SCHIFINO-WITTMANN M.T., MIOTTO
S.T.S., BATTISTIN A. and VALLS J.F.M., 2000b –
Determinação do número de cromossomos e comportamento meiótico em espécies de Adesmia DC.
(Leguminosae). Genetics and Molecular Biology
23, (Suppl): 380.
Received April 26, 2002; accepted June 26, 2002