BRAZILIAN JOURNAL OF PLANT PHYSIOLOGY

The official journal of the REVIEW

Brazilian Society of Plant Physiology

Seed germination in Cerrado species

Lilian B. P. Zaidan1,* and Rosana C. Carreira2

1
Seção de Fisiologia e Bioquímica de Plantas, Instituto de Botânica, Caixa Postal 3005, 01061-970 São Paulo, SP,
Brasil. 2Universidade Cruzeiro do Sul (UNICSUL), Av. Dr. Ussiel Cirilo, 225, CBS, 08060-070 São Paulo, SP, Brasil.
*Corresponding author: lilianzaidan@uol.com.br
Received: 26 August 2008; Accepted: 31 October 2008

The aim of this review is to comment on the available data about germination of seeds from herb, shrub and tree species of the
Cerrado after the publication of the review written by Felippe and Silva in 1984. Studies on seed germination of herbaceous
species focused mainly on the responses of seeds to light, different ranges of temperature and storage in the soil. The majority
of seeds from herb species germinate between 20°C and 30°C, and are photoblastic. Alternate temperatures favored
germination in some seeds, but changed light sensitivity of the achenes of Bidens gardneri. Seeds of most of the shrub
species of Melastomataceae are positive photoblastic; among the Velloziaceae, germination in the dark was observed in some
species. Other shrub species show dormancy caused by impermeability of the seed tegument, as described for seeds of some
species of Bauhinia. Their dormancy is broken by chemical scarification using sulphuric acid. Seeds of Heteropterys
pteropetala are sensitive to very high temperatures, similar to those registered during burnings. Some species were
considered to have allelopathic effects and could inhibit the germination of seeds of other species and the establishment of
plantlets. Seeds of most tree species do not require light to germinate and the focus of the studies were on methods of
dormancy breaking. Sulphuric acid and incisions in the tegument proved to be the most efficient methods to break dormancy.
In general, the studies are limited to three major aspects: responses to light, effects of different temperatures and dormancy
breaking. More studies are necessary to understand the physiological and biochemical aspects of reserve compounds and
their mobilization during germination, as well as the effects of fire in these seeds.
Key words: dormancy, photoblastism, soil seed bank, temperature

Germinação de sementes de espécies de Cerrado: O objetivo desta revisão é comentar dados disponíveis sobre a germinação
de sementes de espécies herbáceas, arbustivas e arbóreas do Cerrado, desde a publicação da revisão feita por Felippe e Silva
na década de 1980. Estudos de germinação de sementes de espécies herbáceas do Cerrado enfatizam as respostas das
sementes à luz, temperatura e armazenamento no solo. A maioria dessas sementes germina entre 20ºC e 30ºC e são
fotoblásticas. Temperaturas alternadas nem sempre favoreceram a germinação das sementes, no entanto, alteraram a
sensibilidade à luz em aquênios de Bidens gardneri. A maioria das sementes das espécies arbustivas de Melastomataceae
estudadas apresentaram fotoblastismo positivo; entre as Velloziaceae, foi observada germinação no escuro em algumas
espécies. Sementes de algumas espécies arbustivas possuem dormência causada pela impermeabilidade do tegumento,
quebrada com imersão em ácido sulfúrico, como no gênero Bauhinia. Sementes de Heteropterys pteropetala são sensíveis
a temperaturas altas, similares àquelas registradas durante as queimadas. Plantas de algumas espécies têm efeitos alelopáticos
e podem inibir a germinação de sementes de outras espécies e o estabelecimento de plântulas. A maioria das sementes de
espécies arbóreas não requer luz para germinar e grande parte dos estudos teve como objetivo testar métodos de quebra de
dormência. Os métodos mais eficazes para a quebra dos diferentes tipos de dormência foram passagem por ácido sulfúrico e
incisões no tegumento das sementes. Em termos gerais, os estudos realizados limitaram-se a três aspectos: fotoblastismo,
temperatura e quebra de dormência. Ainda há grande falta de dados para o entendimento dos aspectos fisiológicos e
bioquímicos da composição e mobilização das reservas das sementes e do efeito do fogo na germinação dessas sementes.
Palavras-chave: banco de semente do solo, dormência, fotoblastismo, temperatura

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168 L. B. P. ZAIDAN and R. C. CARREIRA

INTRODUCTION during the day-time. They are also growing in dry and
shallow soils which are poor in mineral and organic
The Cerrado, the second largest biome in Brazil after
nutrients (Giulietti et al., 1987). In recent years, special
the Amazon forest, hosts a high biodiversity and is listed
attention was given to the germination of seeds of plants
as one of the 34 most endangered ecosystems (hotspots)
of the Serra do Cipó. Most of the plants that had their
of the world (Mittermeier et al., 1999). The cerrado
seed germination investigated are herbs or shrubs and
vegetation is constituted basically by two distinct
many of them are considered as endangered species.
components, a tree/shrub and an herbaceous/subshrub
They belong mostly to the Asteraceae, Eriocaulaceae,
stratum, this latter one comprising more than double the
Fabaceae, Lythraceae, Melastomataceae, Velloziaceae
number of tree species. Some authors claim that herb
and Xyridaceae.
species may account for approximately 80 % of the flora
The aim of this review is to comment on the results
(Mantovani and Martins, 1988; Coutinho, 2002).
obtained since the publication of Felippe and Silva’s review
The woody component presents root systems that
(1984). We did not include in this review data about the
allow the plants to reach deep soil layers with permanent
germination of seeds from species of Riverine Forests,
water availability while the herbaceous component is
Deciduous and Semideciduous Forests. Although there is a
formed by perennial species with well developed
considerable amount of important results in PhD theses and
underground organs, such as rhizomes, bulbs, xylopodia
of data presented in scientific meetings, we decided to
and tuberous roots that guarantee their survival during
consider only the studies effectively converted in scientific
the dry period and occasional fire events. These
papers. It must be mentioned the contribution of the “Rede
underground reserve organs act also as reproductive
de Sementes do Cerrado” (Salomão et al., 2003), with the
structures and for many years plant reproduction in the
publication of a valuable guide for all those interested in the
cerrado was considered to be only vegetative, since seed
germination of seeds of this biome (www.rededesementes
germination was thought to occur only under unique
docerrado. com.br). To facilitate for the reader, the species
circumstances (Ferri, 1960). Later on, it was shown that
that had their seeds investigated were separated into
the germination of seeds in the Cerrado was a relatively
herbaceous, shrub/subshrub and tree species, as defined
common feature and seedlings of 50 woody species were
by Radford et al. (1976).
found growing in the cerrados (Labouriau et al., 1963,
1964; Válio and Moraes, 1966). The early studies on seed
germination of Cerrado species were reviewed by Felippe HERBACEOUS SPECIES
and Silva (1984). Since this review, several additional The Poaceae constitutes the most representative
studies appeared in the current literature, most of them family in the Cerrado. Nevertheless, there is a lack of
describing seed requirements for germination and information about the germination of seeds of native
storage conditions. At the end of the 1990’s studies on cerrado grasses, probably due to their successful
cerrado soil seed banks attracted the interest of some vegetative propagation. So far, the Asteraceae are the
investigators and a few data were published (Sassaki et best represented plant group in studies on seed
al., 1999a, b, c; Pereira-Diniz and Ranal, 2006; Araújo and germination of herbaceous species (Table 1), followed by
Cardoso, 2006, 2007). the Xyridaceae, Eriocaulaceae and Velloziaceae. Some
Within the Cerrado biome, the vegetation known as species of these three latter families are endemic to the
“campos rupestres” is considered to host a number of Serra do Cipó (Minas Gerais State, Brazil). The seeds
endemic and threatened species (Lara and Fernandes, usually require light to germinate, except for Vellozia
1996). Studies carried out in the Serra do Cipó (Minas epidendroides Mart. ex Schult. & Schult., described as
Gerais State, Brazil) allowed us to conclude that the negatively photoblastic, according to Garcia et al. (2007).
plants from this vegetation are well adapted to the The achenes of Bidens gardneri Baker (Asteraceae)
environmental stress conditions they are subjected to, can be separated in short, medium and long achenes. The
such as high exposure to sunlight and to large variations short achenes are located in the margins of the capitulum
in temperature not only during summer/winter, but also and show lower germinability when compared to long

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 GERMINATION ECOPHYSIOLOGY OF CERRADO SEEDS 169

Table 1. Light/dark requirement, type of dormancy and/or characteristics of seeds of herbaceous species of the Cerrado.

Species Family Light/Dark Type of dormancyand/or Reference

requirement other characteristics
Bidens gardneri Asteraceae light germination % is Felippe (1990a);
Baker (nonphotoblastic higher in longer Sassaki et al. (1999a);
under alternate achenes; high % Sassaki et al. (1999b)
temperatures) of germination in
light is maintained
during 6 month
storage at 4°C and
decreases to 60%
after 9 month
storage
Dychia tuberosa (Vell.) Bromeliaceae indifferent optimum temperature Vieira et al. (2007)
Beer between 30°C and 35°C
Eremanthus Asteraceae indifferent campo rupestre; Velten and Garcia (2005)
elaeagnus (C. Martius germination % is low,
ex DC.) Schultz-Bip; E. due mainly to absence
glomerulatus Less.; E. of embryos; alternate
incanus (Less.) Less. temperature did not
promote germination
Paepalanthus Eriocaulaceae light campo rupestre; Sá e Carvalho and
specious Koern. thermo-resistant Ribeiro (1994)
(80ºC for 30min)
Psychotria Rubiaceae light (newly low germination % Sassaki et al. (1999a)
barbiflora DC. collected )/ when stored at 4°C
indifferent (after and in cerrado soil
storage in soil)
Schizocentron Melastomataceae light germination % is Carreira and Zaidan
elegans Meissn. higher between 20ºC (2007)
and 30ºC
Syngonanthus elegans Eriocaulaceae light campo rupestre; Oliveira and Garcia
(Bong.) Ruhland; S. alternate temperatures (2005)
elegantulus Ruhland; favour germination;
S. venustus Silveira;
S. nitens (Bong.) acidity and hypoxia do Schmidt et al. (2008)
Ruhland not affect germination
Tibouchina gracilis Melastomataceae light germination % is higher Carreira and Zaidan
(Bonpl.) Cogn. under light and (2007)
between 20ºC and 30ºC
Vellozia glandulifera Velloziaceae light campo rupestre; seeds Garcia and Diniz (2003)
Goethart & Henrard; must be exposed to high
V. variabilis Mart light intensities
ex Schult. & Schult
Vellozia epidendroides Velloziaceae indifferent campo rupestre; may Garcia et al. (2007)
Mart ex Schult. form soil seed banks;
& Schult. germinates at different
temperatures and light
conditions
Vernonia cognata Asteraceae indifferent achenes are viable for Cesarino and Zaidan
Less. over 18 months if stored (1998)
at 4°C but lose viability

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170 L. B. P. ZAIDAN and R. C. CARREIRA

after 10 month storage at

room temperature
V. herbacea (Vell.) Asteraceae light? (very low % germination of newly Sassaki et al. (1999a)
Rusby of germination) collected seeds was 11%
in light and 3% in the dark
after 25 days; only 15% of
the achenes have embryos
(tetrazolium test)
Xyris cipoensis Xyridaceae light campo rupestre; Abreu and Garcia
Smith & Downs; X. alternate temperatures (2005)
longiscarpa Alb. did not favour germination
Nilsson; X. platystachia
Alb. Nilsson; X.
trachyphilla Mart.

achenes. Germination is high under continuous light and Ruhland and S. venustus Silveira (Oliveira and Garcia,
low in darkness but increases during the period of 2005), but had no effect on the germination of seeds of
storage (Felippe, 1990a). Achenes stored for up to six some species of Eremanthus (Velten and Garcia, 2005),
months at 4°C showed light sensivity, but after nine Schizocentron elegans Meissn. and Tibouchina gracilis
months storage, the photoblastic response is lost for the (Bonpl.) Cogn. (Carreira and Zaidan, 2007).
shortest and longest achenes (Sassaki et al., 1999b). Another effect of alternate temperatures is to
Alternate temperatures during storage of achenes with substitute the positive effects of light on the germination
high moisture content followed by alternate temperatures of photoblastic seeds, as seen in B. gaardneri, although
during germination change the light sensitivity of the this effect was not observed in seeds of some species of
achenes (Rondon et al., 2001). According to the authors, Xyris (Abreu and Garcia, 2005) and of some
this could explain the germination of the achenes during Melastomataceae, as seen in S. elegans and T. gracilis
storage in the soil. (Carreira and Zaidan, 2007). The period of storage, both in
In general, studies on seed germination focus on the the soil and in refrigerator at 4°C, affected the
responses of seeds to light, different ranges of photoblastism of the seeds of Psychotria barbiflora DC.
temperature, here included alternate temperatures, and In this species, newly collected seeds are positively
storage in the soil. Seeds of the Eriocaulaceae photoblastic but become indifferent to light during
Paepalanthus speciosus Koern. (Sá e Carvalho and storage (Sassaki et al., 1999a).
Ribeiro, 1994) and of the bromeliad Dichya tuberosa Seeds (achenes) of Eremanthus ssp. and Vernonia
(Vell.) Beer (Vieira et al., 2007) germinate more rapidly at herbacea (Vell.) Rusby show low germinability. The
temperatures higher than 30°C. This could increase the production of achenes by species of Eremanthus is high,
capacity of the species to survive after fire. At lower but many of them do not complete their development or
temperatures, germination is very heterogeneous, suffer predation by insects. In apparently viable achenes
leading to the formation of seedlings and plantlets at of E. elaeagnus E. glomerulatus, about 91% and 70%,
different times; plant growth and flowering will occur at respectively, had no embryos (Velten and Garcia, 2005). In
different times causing a lost in the synchronisation of V. herbacea, 15% of the achenes did not have viable
this process, an important condition for sexual embryos according to the tetrazolium test (Sassaki et al.,
reproduction. Nevertheless, the majority of seeds from 1999a).
herb species of the Cerrado and “Campos Rupestres” The floral scapes of Syngonanthus nitens (Bong.)
germinate between 20°C and 30°C, as already mentioned Ruhland (Eriocaulaceae) are used in handcrafts, being an
by Felippe and Silva (1984) for Cerrado seeds. Alternate important source of income in the Jalapão region (State of
temperatures favored germination in seeds of Tocantins). For this reason, a study on the conditions
Syngonanthus elegantulus Ruhland, S. elegans (Bong.) that regulate the germination of these seeds was

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 GERMINATION ECOPHYSIOLOGY OF CERRADO SEEDS 171

conducted (Schimidt et al., 2008). The authors verified 15°C and 35°C had a negative effect on germination and
that seed germinability is dependent on the time of seed this corroborates the fact that temperatures between 20°C
production and seed dispersal. Seeds produced in and 30°C are favorable to start the germination process
September/October had higher germinability. The seeds (Felippe and Silva, 1984; Melo et al., 1998). In relation to
require light to germinate and can be stored at -20ºC. 25ºC, application of alternate temperatures did not
These data give important information to support the abbreviate the time to start germination nor the
economic use of the species. germinability of the seeds of the Melastomataceae
The African grasses, Melinis minutiflora Beauv and species tested.
Brachiaria decumbens (Nees) Stapf. have a great The distribution of plants of Melastomataceae
competitive ability over cerrado herbaceous species and species in areas of Cerrado and “Campos Rupestres”
are considered the most threatening invasive plants in could be influenced by the quality and quantity of light
the biome Actually, these grasses occur in virtually every radiation that reaches the seeds. In a secondary way,
cerrado fragment, outcompeting native herbs (Klink, temperature could affect seed germination, probably
1996; Pivello et al., 1999). This fact raised investigations acting on the speed of the germination process. The
about the possibility of the plants to have allelopathic Melastomataceae produce large amounts of small seeds
effects. This was recently confirmed in the study carried with high longevity, remaining viable during years, both
out by Barbosa et al. (2008). The authors demonstrated in laboratory conditions and buried in the soil, isolated or
that aqueous leachates of leaves and seeds of B. within the fruits (Carreira and Zaidan, 2003). In the
decumbens were able to inhibit the germination of seeds absence of light they do not germinate and may remain
of test species (Lactuca sativa L.) and of other invasive dormant in the soil for long periods thus constituting soil
plants (M. minutiflora and Phalaris canarinensis L.). seed banks. In fact, in the studies of soil seed banks in
the Cerrado, the most common seeds were always those
of the Melastomataceae (Sassaki et al., 1999a). Plants
SHRUB/SUBSHRUB SPECIES
belonging to the Rubiaceae (Palicourea marcagravii St.
Studies on the germination of seeds of shrub and Hil., Psychotria hoffmanseggiana (Wild. Ex Roem. &
subshrub species from the Cerrado are focused mainly on Schutz.) Mull. Arg. and P. vellosiana Benth.) also
the sensitivity of seeds to light and on optimum or contribute to form soil seed banks (Araújo and Cardoso,
cardinal temperatures. Effects of light and temperature 2007). According to the authors, artificial storage in the
are the most common methods to evaluate germination, in soil favored the survival of the seeds and contributed to
terms of germinability of seeds and the mean speed of maintain their longevity. Production of seeds that can
germination. The germination tests are usually keep their viability for long periods in the soil is important
conducted at temperatures in the range from 15°C to 40°C to individuals and to a population, since it allows the
at 5°C intervals, and sometimes under alternate colonization of a great variety of habitats (Garcia et al.,
temperatures (30°C/20°C) under continuous light/ 2007).
darkness. Some shrub species show dormancy caused by
Seeds of all species belonging to the impermeability of the tegument of the seeds. This seems
Melastomataceae that had been investigated by Carreira to be the case of seeds of some species of Bauhinia. In
and Zaidan (2007) are positively photoblastic and no these seeds, dormacy is broken with chemical
germination in darkness was registered. Nevertheless, scarification using sulphuric acid (Pereira, 1992; Alves et
Ranieri et al. (2003) and Silveira et al. (2004) observed al., 2000). Another common species of the Cerrado is
some germination in the dark. Among the Velloziaceae, Solanum lycocarpum St. Hil., common name “fruta-do-
some germination in the dark was also observed (Garcia et lobo”. The fruit is a basic component of the diet of the
al., 2007), as shown in Table 2. “lobo-guará” (Chrysocyum brachyurus Illiger) and thus,
Higher germinability was achieved at 20°C and 30°C seeds are frequently found in the faeces of this animal
in seeds of the Melastomataceae of the Cerrado and the (Dietz, 1984). Although these seeds pass through the
Velloziaceae of the “Campos Rupestres”. Temperatures of digestive tract of the animal, thus suffering acid

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172 L. B. P. ZAIDAN and R. C. CARREIRA

Table 2. Light/dark requirement, type of dormancy and/or characteristics of seeds of shrub/subshrub species of the
Cerrado.

Species Family Light/Dark Type of dormancy and/or Reference

requirement other characteristics
Actinocladum Poaceae light germination % is higher Felippe and Filgueiras
verticilatum (Ness) at 25ºC (1986)
McClure ex Soderstrom
Andira humilis Mart. Fabaceae ? (unknown) allelopathic potential Periotto et al. (2004)
ex Benth
Baccharis Asteraceae indifferent campo rupestre; higher Gomes and Fernandes
dracunculifolia D.C. EVIs in achenes (2002)
germinated at 15°C and
20ºC under light and 15ºC
in the dark
Bauhinia forficata Fabaceae ? (unknown) tegument impermeability; Pereira (1992)
Link. the best treatment: 30ºC
and vermiculite
B. monandra Britt.; Fabaceae light tegument impermeability; Alves et al. (2000)
B. ungulata L. dormancy break:
scarification with
sulphuric acid
Byrsonima intermedia Malpighiaceae ? (experiment in vitro germination; Nogueira et al. (2004)
A. Juss. under 16 h best culture media: MS
photoperiod) and WPM 50%, without
sucrose
Campomanesia Myrtaceae ? (unknown) recalcitrant Melchior et al. (2006)
adamantium Camb.
C. pubescens (DC.) Myrtaceae indifferent ? (unknown); shorter Arrigoni-Blank et al.
Berg. time to germinate at 25ºC (1997)
Diplusodon virgatus Lythraceae light storage at 5ºC; seeds Cesarino et al. (1998)
Pohl. remain viable after 12
months
Heteropterys Malpighiaceae indifferent seeds affected by Schmidt et al. (2005)
pteropetala early-fire
(Adr. Juss.)
Lavoisiera cordata Melastomataceae ? (experiment campo rupestrehighest Ranieri et al. (2003)
Cogn. under 12 h radicle emergency rate at
photoperiod) 20ºC and above
L. francavillana Cogn. Melastomataceae ? (experiment campo rupestrehighest Ranieri et al. (2003)
under 12 h radicle emergency rate at
photoperiod) 25ºC and above;
Marcetia taxifolia (A. Melastomataceae light campo rupestre higher Silveira et al. (2004)
St.-Hil.) DC. germination % at 15°C
and 20ºC
Miconia albicans (Sw.) Melastomataceae light allelopathic potential; Gorla and Perez (1997);
Triana forms soil seed banks; Carreira and Zaidan
germination is higher (2007)
under light and between
20ºC and 30ºC
M. langsdorffii Cogn.; Melastomataceae light germination is higher Carreira and Zaidan
M. stenostachya Schr. under light and between (2007)
& Mart. ex. DC. 20ºC and 30ºC

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 GERMINATION ECOPHYSIOLOGY OF CERRADO SEEDS 173

Palicourea Rubiaceae light potentiality to form soil Araújo and Cardoso

marcagravii st. Hil. seed banks; soil storage (2007)
favoured seed survival
and germination
Platycyamus regnelli Fabaceae indifferent ? (unknown); germination Scalon et al. (1993)
Benth. is higher at 25°C and 30ºC
and in roled paper and
between paper layers
Psychotria Rubiaceae light potentiality to form soil Araújo and Cardoso
hoffmansegiana (Wild. seed banks; soil storage (2007)
ex. Roem. & Schult.) favoured seed survival
Mull. Arg. and germination
P. vellosiana Benth. Rubiaceae light potentiality to form soil Araújo and Cardoso
seed banks; seeds show (2006)
true dormancy and/or
required an extended time
to germinate
Solanum lycocarpum Solanaceae ? (unknown) tegument impermeability; Monteiro and Ramos
St. Hil. allelopathic potential (1997); Oliveira et al.
(2004); Aires et al.
(2005)
Stylosanthes Fabaceae indifferent germination is higher Silva and Felippe (1986)
macrocephala M.B. between 20ºC and 30ºC
Ferri & Souza Costa
Vellozia leptopetala Velloziaceae light campo rupestre Garcia et al. (2007)
Goeth. et Henr. potentiality to form soil
seed banks; germination
in different temperatures
and light conditions
V. gigantea N.L. Velloziaceae light campo rupestreseeds Garcia and Diniz (2003)
Menezes & Mello-Silva submitted to high light
intensities
Zeyhera montana Mart. Bignoniaceae ? (experiment embrionic stretching and Dousseau et al. (2007)
under 12 h possible presence of
photoperiod) inhibitors

scarification, differences in the germination of seeds exposed to temperatures of 60°C during 40 minutes, 80°C
collected in faeces or in mature fruits were not detected. In during 10 minutes and 100°C during 2 minutes. None of
both group of seeds, the percentage of germination was these treatments interfered in the viability (≥ 80%) nor in the
70%. The author observed that when the animal bites the germination of the seeds (≥ 70%). Treatments of 100°C
fruit it causes damages to the seeds so that the percentage during 5 or 10 minutes and of 200°C during 1 minute reduced
of seed germination should be higher than 70% (Monteiro the germinability respectively in 50%, 90% and 100%. In
and Ramos, 1997). contrast, late burnings (area of the RECOR submitted to
Some seeds are sensitive to very high temperatures, biennial fires towards the end of the dry period in
similar to those registered during burnings. Seeds of September) had positive effects and caused increases in the
Heteropterys pteropetala (Adr. Juss.), Malpighiaceae, number of individuals recruited (Schmidt et al., 2005).
collected in a natural area in Brasília (“Reserva Ecológica do Oliveira (1998) observed that fire may interfere on the
IBGE” – RECOR, DF) showed to be sensitive to early structure and the composition of species in a community
burnings (areas of the RECOR submitted to biennial fires in through its effects on sexual reproduction. Nevertheless,
June), as reported by Schmidt et al. (2005). The seeds were few studies have been conducted using high temperatures

Braz. J. Plant Physiol., 20(3):167-181, 2008

174 L. B. P. ZAIDAN and R. C. CARREIRA

similar to those found during the passage of fire. germination (Table 3). Dormancy is broken when the seeds
Miconia albicans (Sw.) Triana (Gorla and Perez, 1997; are sunk in gibberellic acid (Bernardes et al., 2007). Seeds of
Carreira and Zaidan, 2007), Andira humilis Mart. ex Benth Dipteryx alata Vog. need a period of post maturation inside
(Periotto et al., 2004), S. lycocarpum (Oliveira et al., 2004, the fruit to achieve high germinability (Corrêa et al., 2000).
Aires et al., 2005) were considered to have allelopathic Seeds of Eugenia calycina Cambess. and Tabebuia
effects. From these, only the seeds of M. albicans were heptaphylla (Vell.) Toledo show low viability after
experimentally tested and showed to be affected by their harvest. These species are cultivated for fruit production
own extracts (Carreira and Zaidan, 2007). These authors and ornamental purposes, respectively. For this reason
mentioned that field observations point to the experiments aiming to obtain homegeneous plantlets in
hypothesis of an allelopathic effect on other plant the field were carried out to test differents substrates
species, considering that plants of the same species form (Büllow et al., 1994). Seeds of T. heptaphylla showed
homogenous groups and could inhibit the germination of higher germinability int the presence of clay cerrado soils
certain seeds and the establishment of others. Dousseau with or without organic matter (Bocchese et al., 2008).
et al. (2007) observed a kind of embryonary dormancy in Some seeds have peculiar requirements to germinate.
seeds of Zeyera montana Mart. and discussed the This is the case of seeds of Peltophorum dubium Taub.
possible presence of germination inhibitors, although the and Pterogyne nitens Tul. Seeds of P. dubium are
authors failed to isolate this inhibiting agent. temperature-resistent: exposition for 24 hours at 45ºC is
sufficient to break the mechanical dormancy imposed by
the tegument (Perez et al., 1998). Gibberellic acid allows
TREE SPECIES
an increase in the maximum tolerance to water stress
The seeds of most of the species seem to have no light especially in the presence of putrescin and espermidin,
requirement to germinate, i.e. they do not show growth regulating substances that modulate some
photoblastic response. Nevertheless, one must consider biological process such as cell division, responses to
that in some studies the germination tests were stressing factors and developmental processes (Botelho
conducted directly in the soil and the information about and Perez, 2001). Seeds of P. nitens are resistant to water
the light conditions during the experimental period was stress and to several salt solutions (Nassif and Perez,
omitted. For this reason studies on seed germination of 1997a). The seeds are tolerant to dryness between -2.4
tree species of the Cerrado focus on methods of and -2.6 MPa (manitol) and -1.0 and -1.2 MPa (PEG 6000).
dormancy breaking and tegument impermability. Using CaCl 2 e KCl solutions, the maximum limit of
Seeds of most savanna trees are dormant and physical tolerance is between -1.6 and -1.8 MPa and for NaCl
dormancy is the most common type (Baskin and Baskin, solutions, between -2.0 and -2.2 MPa.
2001). The impermeability of the tegument to water and Felippe (1990b) and Godoy and Felippe (1992) studied
oxygen is a kind of dormancy rather common in large seeds, the imbibition and the germination of the seeds of Qualea
and especially in those of the Fabaceae. This type of grandiflora Mart. and Q. cordata Spreng., respectively.
dormancy can be broken by scarification resulting in the The authors verified that both species have light colour
rupture or weakness of the tegument, allowing the entrance and dark colour seeds the latter ones being more frequent.
of water and oxygen and thus leading to the starting of the The light colour seeds imbibe more quickly than the dark
germination process. There are many methods to surpass colour seeds and their germinability is around 80%. The
this type of dormancy like mechanical scarification using tetrazolium test showed that only 2% of the dark colour
abrasive materials, incisions or damages in the tegument, seeds of Q. grandiflora are viable (Felippe, 1990b).
stratification or exposition to high temperatures and
chemical scarification by strong acids. Among these
methods, sulphuric acid and incisions in the tegument
CONCLUDING REMARKS
proved to be the most efficient. The review of the more recent literature on seed
The seeds of Annona crassiflora Mart. exhibit germination of Cerrado species shows that the studies
embrionary dormancy that causes desuniformity in are limited to tests on seed responses to light (mainly

Braz. J. Plant Physiol., 20(3):167-181, 2008

 GERMINATION ECOPHYSIOLOGY OF CERRADO SEEDS 175

Table 3. Light/dark requirement, type of dormancy and/or characteristics of seeds of herbaceous species of the Cerrado.

Species Family Light/Dark Type of dormancy and/or Reference

requirement other characteristics
Annona crassiflora Annonaceae ? (unknown) embrionary; GA3 1000 Bernardes et al. (2007)
Mart. ppm was efficient to
break dormancy
Astronium urundeuva Anacardiaceae ? (experiment seeds are ortodox and Medeiros and Cavallari
(Fr. All.) Engl. under 12 h germination is higher at (1992)
photoperiod) 20ºC
Bowdichia virgilioides Fabaceae ? (unknown) tegument impermeability; Smiderle and Sousa
Kunth best method for (2003); Albuquerque et
dormancy break: al. (2007)
scarification with
sulphuric acid
Caryocar brasiliensis Caryocaraceae ? (unknown) undifferentiated Sá e Carvalho et al.
Camb. embryos; allelopathic (1994); Melo and
potential Gonçalves (2001)
Cassia excelsa Schrad Fabaceae ? (unknown) tegument impermeability; Jeller and Perez (1999);
dormancy break with Jeller et al. (2003)
sulphuric acid; highest
germinability achieved
after conditioning with
distilled water
Copaifera langsdorffii Fabaceae ? (unknown) tegument impermeability; Perez and Prado (1993)
Desf. seeds immersed in ether
and germination
inhibited by dry hot
application and boiled
treatment
Dalbergia miscolobium Papilionaceae indifferent germination is concluded Arasaki and Felippe
Benth. in 6 days (1987)
(sin. D. violacea)
Dipteryx alata Vog. Fabaceae ? (unknown) post-maturation; no Corrêa et al. (2000)
variation in the % of
plant emersion and
emersion velocity index
Enterolobium Fabaceae ? (germination in tegument impermeability; Eira et al. (1993);
contorsiliquum (Vell.) the soil) better method for Monteiro and Ramos
Morong. dormancy break is (1997)Lima et al. (1997)
scarification with
sandpaper;Tmax
between 40.9ºC and
42.4ºC
Eugenia dysenterica Myrtaceae indifferent better results with sandy Nietsche et al. (2004);
Mart. ex. DC. and clay; larger seeds Duarte et al. (2006);
from apparently more Martinotto et al. (2007)
vigorous fruits
E. calycina Cambess. Myrtaceae ? (unknown) ? (unknown); loses Bülow et al. (1994)
viability quickly
Kielmeyera coriacea Guttiferae ? (unknown) higher viability when Botelho and Carneiro
Mart. kept in cold conditions (1992)
Miconia rubiginosa Melastomataceae light germination % is higher Carreira and Zaidan

Braz. J. Plant Physiol., 20(3):167-181, 2008

176 L. B. P. ZAIDAN and R. C. CARREIRA

(Bonpl.) DC. under light and between (2007)

20ºC and 30ºC
Myracrodruon Anacardiaceae indifferent ? (unknown); Medeiros et al. (2000);
urundeuva Allemão polyembriony; Silva et al. (2002);
stratificationduring six Dorneles et al. (2005);
days; germination % is Pacheco et al. (2006)
higher in vermiculite
and coconut fiber
substrates
Ouratea spectabilis Ochnaceae ? (unknown) allelopathic potential Silva et al. (2006)
(mart.) Engl.
Peltophorum dubium Fabaceae indifferent tegument impermeability; Perez et al. (1998; 1999);
Taub. thermo-resistant; GA3 De Fiore and Perez
extended the maximal (2000); Botelho et al.
tolerance limit to water (2001)
stress
Platypodium elegans Fabaceae ? (unknown) tegument Pacheco et al. (2007)
Vog. impermeability;
longitudinal cuts
promote higher
indices of germination
speed
Pouteria ramiflora Sapotaceae ? (unknown) allelopathic potential Silva et al. (2006)
(Mart.) Radlk.
Pterogyne nitens Tul. Fabaceae ? (unknown) tegument impermeability; Nassif and Perez
best method for (1997a; 1997b; 2000)
dormancy break is
scarification with
sulphuric acid; tolerance
to salt solutions
Qualea grandiflora Vochysiaceae indifferent dark seeds with no Felippe (1990b); Silva
Mart. embryos; allelopathic et al. (2006)
potential
Q. cordata Spreng. Vochysiaceae ? (unknown) dark seeds with no Godoy and Felippe
embryos (1992)
Senna macranthera Fabaceae ? (unknown) tegument impermeability; Eschiapati-Ferreira and
(Collad.) Irwin et. Barn. immersion in sulphuric Perez (1997)
acid
Shiphoneugena Myrtaceae ? (germination in ? (unknown); depulped Monteiro and Ramos
densiflora Berg. the soil) fruits germinated twice (1997)
as much as intact fruits
Stryphnodendron Fabaceae ? (unknown) allelopathic potential Barreiro et al. (2005);
adstringens (Mart.) Silva et al. (2006)
Coville
Tabebuia Bignoniaceae ? (experiment low viability; higher seed Bocchese et al. (2008)
heptaphylla (Vell.) under a shaded germination rates in clay
Toledo house) + organicmaterial and
clay soils
Talauma ovata St. Hil. Magnoliaceae ? (germination in ? (unknown); germination Monteiro and Ramos
the soil) is less than 50% with or (1997)
without aril
Tapura amazonica Dichapetalaceae ? (germination in ? (unknown); mechanical Monteiro and Ramos
Poepp. and Endl. the soil) seed coat scarification (1997)

Braz. J. Plant Physiol., 20(3):167-181, 2008

 GERMINATION ECOPHYSIOLOGY OF CERRADO SEEDS 177

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