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... Growth and frutactan contents ...

89

EFFECT O MINERAL NUTRIENTS ON GROWTH AND


FRUCTAN CONTENTS IN PLANTS OF Vernonia
1
herbacea
Patricia G. Texeira 2 , M. Angela Machado de Carvalho 3 , Lilian B.P.
Zaidan 3 and Aldo L. Klein 4

Seçcão de Fisiologia e Bioquímica, Instituto de Botânica, CP 4005, São


Paulo, SP, 01061-970, Brazil.
ABSTRACT- Plants of Vernonia herbacea (Vell.)
Rusby (Asteraceae) were grown under cerrado EFEITO DE NUTRIENTES MINERAIS NO
covered soil or washed solution, solution without
nitrogen, solution without phosphorus, and distilled CRESCIMENTO E NO CONTEÚDO DE
water. In cerrado soil, although shoot dry mass and FRUTANOS EM PLANTAS DE Vernonia
leaf area were higher when complete solution was
supplied to plants, fructan contents were significantly
herbacea
lower in the underground organs, suggesting the RESUMO- Plantas de Vernonia herbacea (Vell.)
translocation of these sugars to the growing tissues in Rusby (Asteraceae) foram cultivadas em terra sob
the plant tops. the lack o nitrogen did not promote cerrado ou areia lavada, em casa-de-vegetação
growth of rhizophores in sand or growth of plants in adicionando-se: solução de Hoagland completa,
either substrate and the plants accumulated small solução sem nitrogênio, solução sem fósforo e água
amounts of fructans, specially fructo-polysaccharides. destilada. Em solo sob cerrado, embora a massa seca
It is suggested that nitrogen is important to the growth da parte aérea e a área foliar tenham sido maiores nas
of this plant. Vegetative growth and fructan plantas que receberam solução de Hoagland
accumulation were not specially affected by the completa, os teores de frutano foram
absence of phosphorus. Neither nutrient treatments significativamente menores, sugerindo a translocação
nor substrate influenced the distribution pattern of the desses açúcares para os tecidos em crescimento na
components of the homologous series of fructans. The parte aérea das plantas. A ausência de nitrogênio não
data here obtained indicate that nutrient condition is promoveu o crescimento dos rizóforos, em areia, nem
one of the factors defining growth and accumulation o crescimento das plantas, em ambos substratos, e as
patterns in this species. Plants of V. herbacea naturally plantas acumularam pequenas quantidades de
growing in the cerrado seem to have developed frutanos, especialmente polifrutanos. Sugere-se que o
adaptive strategies that supply them with efficient nitrogênio seja importante para o crescimento desta
mechanisms of carbon partitioning that favors fructan espécie. O crescimento vegetativo e o acúmulo de
accumulation in the rhizophores. frutanos não foram afetados pela ausência de fósforo.
O padrão de distribuição dos componentes da série
Additional index terms: Asteraceae, cerrado, mineral
homóloga de frutanos não foi influenciado pelos
nutrition.
tratamentos de nutrição nem pelos substratos. Os
dados indicaram que a condição nutricional é um dos
fatores que controla o crescimento e o acúmulo de
1 frutanos nesta espécie. Plantas de Vernonia herbacea
Recebido em 30/12/96 e aceito em 06/08/1997.
Financiado pela FAPESP (Proc. 91-3588-8) crescendo naturalmente no cerrado apresentariam
2 estratégias adaptativas bem desenvolvidas que
Bióloga, Bolsista do CNPq permitiriam nos solos oligotróficos do cerrado,
3
Pesq.Científico da Seção de Fisiologia e bioquímica, mecanismos eficientes de partição de carbono que
Instituto de Botânica, CP 4005, São Paulo, SP, 01061, favoreceriam o acúmulo de frutanos em seus
Bolsista CNPq. rizóforos.
4
Prof.Assistente Doutor, Depto de Ciências Biológicas,
Faculdade de Ciências e Letras de Assis, UNESP, CP Termos adicionais para indexação: Asteraceae,
335, Assis, SP, 19800-000. cerrado, nutrição mineral.

R. Bras. Fisiol. Veg., 9(2):89-96, 1997


90

nutrient solution without phosphorus (-P). The control


INTRODUCTION plants received distilled water.

Soils under Brazilian cerrados are, in general, old, Growth measurements


deep (Ferrasols) and leached, with low clay contents, Growth analysis of aerial organs was performed by
low pH and toxic amounts of aluminium (Goodland, measuring stem height and leaf number
1971). Although these characteristics may present approximately every ten days, using the five plants of
some degree of variation among different soils under each treatment. As most of the plants entered
cerrado vegetation, oligotrophy seems to be the only dormancy in winter, all plants with the remaining aerial
feature in common (Arens, 1963). organs had them cut for standardization purposes.
Trees and shrubs show all the characteristics of a Measurements were resumed at the onset of the new
crubby flora, limited by several growth factors (Arens, growth cycle.
1963; Eiten, 1972) other than water availability At the end of the experiment, leaf area and fresh
(Rawitscher, 1942; Ferri, 1944). The herbaceous layer, and dry masses of plant tops and rhizophores were
however, with root systems developing in the upper also obtained. Leaf area was measured using a
part of sandy soils, with low water retention capacity, portable area meter LI_COR-3000 and dry mass was
undergo water stress during the dry season that lasts determined by oven drying at 80oC to constant mass.
from 4 to 7 months (Aámoli et al., 1985). To survive
these conditions, cerrado perennial herbs have Soluble carbohydrate extractions
developed, as a main phenological characteristic, a Fructans were extracted in triplicate according to
long phase of dormancy. While their aerial organs are Pollock & Jones (1979) from samples of fresh
renewed each year, the underground perennial parts rhizophores of 5 plants with 5g each. Tissue was sliced
persists in a latent stage until the onset of a new active and boiled in 80% aqueous ethanol for 3min for
cycle in the rain season (Monasterio & Sarmiento, enzyme denaturation. The mixture was then ground in
1976). As an example, Vernonia herbacea bears an a mortar with a pestle. The homogenate was placed in
underground organ known as rhizophore (Menezes et a water bath at 80oC for 15min and centrifuged at 1000
al., 1979) which accumulates fructans as reserve g for 15min. The residue was re extracted on as above
carbohydrate (Carvalho & Dietrich, 19930. and then submitted twice to water extraction for 30min
It is long known that nitrogen and phosphorus at 60oC. The pooled supernatants were concentrated,
deficiencies promote the accumulation of soluble frozen, thawed and centrifuged at 9000g for 20min at
sugars, mainly fructans (Archbold, 1938; Russel, 5oC. The pellet countaining the high molecular mass
1938), although this effect has not been studied in fructans (fructo-polysaccharides) was resuspended in
cerrado plants. As cerrado soils present low water.
concentrations of nitrogen and phosphorus, the aim of Medium molecular mass fructans present in the
this study was to determine a possible relationship supernatant were precipitated with 3 volumes of
between nutrient conditions and fructan accumulation ethanol and added to the fructo-polysaccharide
in rhizosphores of V. Herbacea plants. fraction. The remaining supernatant constituted the
low molecular mass fructans (fructo-
MATERIAL AND METHODS oligosaccharides).
Plants of Vernonia herbacea (Vell.) Rusby Carbohydrate analyses
(Asteraceae) were obtained from fragments of Free and combined fructose were measured using a
rhizophores of adult plants collected in a preserved ketose-specific modification of the anthrone reaction
cerrado area in Moji Guaçu (22o18’ S, 47o 11’ W) SP, (Jermyn, 1956), using fructose and inulin from
Brazil. The fragments were taken from plants with Helianthus tuberosus as standards.
vigorous underground system, favoring the obtention
of numerous individuals from a single plant, partially Fructo-oligosaccaharides were analyzed by
avoiding eventual genetic variation. The plants were thin-layer chromatography on silica gel plates as
divided in two groups of 20 and cultivated in pots described by Kanaya et al. (1978). Samples were
containing 2.5L of cerrado soil or washed sand, in prepared as described in Carvalho & Dietrich (1993).
glasshouse conditions. Fructans were visualized by spraying with the
ketose-specific urea-phosphoric acid (Wise et al,
Every 10 days, groups of 5 plants in each substrate 1955). Fructo-oligosaccharides of the inulin series
received 50mL of Hoagland nutrient solution from tuber of H. tuberosus were used as reference
(Hoagland & Arnon, 1938) as: complete nutrient material.
solution (H), nutrient solution without nitrogen (-N) and

R. Bras. Fisiol. Veg., 9(2):89-96, 1997


... Growth and frutactan contents ... 91

FIGURE 1- Growth of main shoot of plants of Vernonia FIGURE 2- Number of leaves in plants of Vernonia
herbacea cultivated in cerrado soil (A) and in sand (B), herbacea cultivated in cerrado soil (A) and in sand (B),
under different nutrient treatments. Hoagland nutrient under different nutrient treatments. Hoagland nutrients
p
solution n , nutrient solution without nitrogen (-N) = , solution = n , nutrient solution without nitrogen (-N) =
nutrient solution without phosphorus (-P) = ∆ and ∆, nutrient solution without phosphorus (-P) = p and
control (H2O) = o . The arrow indicates when stems control (H2O) = o . The arrow indicates when stems
were cut. Letters compare different treatments (Tukey were cut. Letters compare different treatments (Tukey
5%), where statistical differences were found. 5%), where statistical differences were found.

After deionization through cation (Dowex-50W, Na+ Data were statistically analyzed by one way and two
form) and (Dowex-1, CL- form) resins, neutral soluble way ANOVA and the least significant differences at 5%
carbohydrates were also analyzed on a 4x25mm level were calculated using the Tukey test.
Corbo Pac PA-1 anion exchange column using a
Dionex DX-300 gradient chromatography system with RESULTS
pulsed amperometric detection (HPAEC/PAD). The
gradient was established according to Shiomi (1993). In cerrado soil, although shoots were longer in -N
Eluent A (150mM NaOH) and eluent B (500mM plants, no significant differences were found among
sodium acetate in 150mM NaOH) were mixed as plants receiving other treatments (Fig. 1A). Plants
follows: 0-1min, 25mM; 1-2min, 25-50mM; 2-14min, cultivated in sand did not present differences in shoot
50-500mM; 14-22min, 500mM; 22-30min, 25mM. The length in the first cycle (Fig. 1B), but differences were
flow rate through the column was 1.0mL min-1. The observed in the second growth cycle when complete
applied PAD potentials for E1 (300ms), E2 (120 ms) Hoagland solution promoted shoot elongation.
and E3 (300ms) were 0.04,0.60 and -0.80V, Leaf number and leaf area increased when plants
respectively, and the output range was 1000nA. were treated with complete Hoagland solution in
HPAEC/PAD elution patterns were compared to those cerrado soil (Fig. 2A, 3A). In sand, larger number of
of reference standards obtained from tubers of H. leaves and leaf area were found when plants received
tuberosus extracted according to Pollock & Jones complete and -P Hoagland solution in the second
(1979). cycle (Fig. 2B, 3B). Statistical analysis showed
Fructo-polysaccharides were analyzed by gel interaction between the two substrates and the
permeation chromatography using a Bio-Gel P-10 treatment using solution without phosphorus for leaf
column as described by Carvalho & Dietrich (1993). area (data not shown).
Relative molecular mass (Mr) was estimated
according to Andrews (1965).

R. Bras. Fisiol. Veg., 9(2):89-96, 1997


92

TABLE 1- Dry mass (g) of aerial organs (S) and soil only plants treated whit complete solution showed
rizophores (R) and S:R ratio of plants of Vernonia this response. Statistical interaction occurred between
herbacea cultivated for 12 months in cerrado soil and substrates and nutrient in -P and control treatments,
in sand, under different nutrient solution without as for dry mass accumulation in shoots.
phosphorous (-P) and control (H2O). Plants growing in sand with water exhibited the
Cerrado soil Sand ration shoot/rhizophore two times lower than in -N and
three times lower than in complete solution and -P
S R S:R S R S:R
treated plants (Table 1). In this last group numerous
H2O 3.0b* 4.5ns. 0.66 1.4b 4.0ab 0.35 roots were observed (data no shown). In soil, the ratio
H 7.3a 5.0ns. 1.46 6.3a 5.4ab 1.16 shoot/rhizophore was higher in plants receiving
-N 3.4b 3.8ns. 0.89 2.1b 3.2b 0.65 complete solution when compared to the other
-P 4.0b 5.4ns. 0.74 6.6a 5.7a 1.15 treatments.

* letters compare values in each column (Tukey 5%) Plants treated with complete Hoagland solution
accumulated lower amounts of total fructans when
n.s. - not significant compared to -P and control plants in cerrado soil.
Table 1 shows that dry mass of rhizophores was not Plants treated with -N solution also presented low
aerated by any of the nutrient solution tested in fructan concentration (Fig. 4A). Fructo-
cerrado soil, while in sand plants grown without polysaccharides were found in higher amounts in
phosphorus produced significantly more dry mass in rhizophores of plants submitted to -P and H2O.
the underground organs than plants cultivated without Addition of nutrient had no effect in contents of
nitrogen. Dry mass accumulation in plant tops was fructo-oligosaccharides in plants grown in sand (Fig.
promote in complete Hoagland solution and -P treated 4B). The variation observed in total fructans reflected
plants grown in sand, but in those cultivate in cerrado the pattern found for fructo-polysaccharides: H2O and
-P treatments favored the accumulation of higher
amounts of these compounds.
TABLE 2- Relative molecular mass (Mr) and degree
of polimerization (DP) of fructo-polysaccharides from
rhizophores of Vernonia herbacea cultivated in
cerrado soil and in sand, under different nutrient
treatments. Hoagland nutrient solution (H), nutrient
solution without nitrogen (-N), nutrient solution
without phosphorous (-P) and contro (H 2O).
Cerrado Soil Sand
Mr DP Mr DP
H2O 2570 16 2962 18
H 2570 16 2570 16
-N 2570 14 2270 14
-P 2570 16 2270 14

Mean degree of polymerization and relative


molecular mass of fructo-polysaccharides varied from
14 to 18 and from 2270 to 2962, respectively (Table 2);
the highest values were detected in control plants
grown in sand.
No statistical interaction between treatments was
found in total fructan amounts. With reference to
fructo-polysaccharides, interaction substrate X
nutrient solution was verified in plants treated with
FIGURE 3- Leaf area of plants of Vernonia herbacea complete solution in both substrates. Interaction was
cultivated in cerrado soil (A) and in sand (B), under also found in the amount of fructo-oligosaccharides in
different nutrients treatments. Hoagland nutrient control and -P treatment, in cerrado soil.
solution, nutrient solution without nitrogen (-N), All members of the homologous series of inulin were
nutrient solution without phosphorus (-P) and control present in all treatments, as revealed by TLC analysis
(H2O).
R. Bras. Fisiol. Veg., 9(2):89-96, 1997
... Growth and frutactan contents ... 93

FIGURE 5- Thin layer chromatography (TLC) of


fructo-oligosaccharides from rhizophores of Vernonia
herbacea cultivated in cerrado soil (A) and in sand (B),
under different nutrient treatments. Hoagland nutrient
solution (H), nutrient solution without nitrogen (-N),
nutrient solution without phosphorus (-P) and control
(H2O). Each lane contained approx. 100µg fructose,
FIGURE 4- Contents of fructo-oligosaccharides, S- sucrose, K-1- Kestose, N- nystose, D.P.- degree of
fructo-polysaccharides and total fructans in polymerization.
rhizophores of Vernonia herbacea cultivated in
cerrado soil (A) and in sand (B), under different chain of events that affects plant growth remains
nutrient treatments. Hoagland nutrient solution , largely speculative (Delhaize & Ryan, 1995).
nutrient solution without nitrogen , nutrient solution Plants of Vernonia herbacea cultivated in cerrado
without phosphorus and control (H2O) = o . soil, showed higher shoot dry mass and leaf area
when Hoagland complete solution was added, total
of fructo-oligosaccharides (Fig. 5). Differences in the fructan contents, however, were significantly lower.
proportions of components can be seen. Fructose was Considering that rhizophore dry mass was similar in all
present in low amounts in most treatments, and was treatments, less fructan accumulation was possibly
more evident in plants receiving complete Hoagland due to a higher sink strength of the aerial organs,
solution in sand and cerrado soil. These results were causing lower translocation of assimilates to the
confirmed by HPLC analyzes as seen in Fig. 6. The rhizophores, in this treatment. Indeed, fructan
proportion of glucose and fructose in plants grown accumulation has been shown to be inhibited by
under complete solution treatment was higher in both favorable growth conditions, as elevated nitrogen
sand and cerrado soil when compared to other nutrient supply associated to high temperatures (Pollock,
treatments. 1986). In plants of V. Herbacea, although the lack of
nitrogen inhibited vegetative growth, fructan
DISCUSSION accumulation was low.
In general, soils under cerrado vegetation are rich in It can be argued that the generally higher growth of
aluminium and plants found in this biome my aerial parts in plants treated with complete solution in
accumulate and/or bear tolerance to this mineral both substrate occurred due to a more intense
element (Goodland, 1971). Thus, the aluminium factor metabolic activity in the rhizophores, where fructans
should be considered whenever adaptive mechanisms are promptly hydrolyzed to sucrose and translocated
to tropical savanna conditions discussed (Sarmiento, to the developing shoot. Evidence of fructan
1984). The presence of aluminium in soils is often mobilization is given by the increase in the proportion
related to a decrease in the availability of P, Ca, Mg, N, of glucose and fructose in these plants (Fig. 6),
K among other elements, leading to mineral deficiency suggesting intense enzymatic activity. the low
symptoms in plants (Goodland, 1971). The most easily amounts of total fructan in plants irrigated with
recognized symptom of aluminium toxicity is the complete Hoagland solution corroborates this
inhibition of root growth. The root apex tissues. But the hypothesis (Fig. 4). In contrast, control plants presents
R. Bras. Fisiol. Veg., 9(2):89-96, 1997
94

FIGURE 6- Qualitative analyses by HPAEC/PAD of fructo-oligosaccharides from rhizophores of Vernonia


herbacea cultivated in cerrado soil (A) and in sand (B), under different nutrient solution without phosphorus (-P)
and control (H2O). G-glucose, F-fructose, S-sucrose, K-1-kestose, N-nystose, numerals 5-7 refer to
oligosaccharides of DP 5-7.

higher fructan accumulation and lower shoot mass. In For major nutrients, plants in general show a lower
sand, control plants presented the highest molecular shoot/root ration when a particular nutrient is deficient
mass fructans which, in this context, suggesting low (Wilson, 1988). Plants of V. herbacea cultivated in
mobilization activity. the values for molecular masses cerrado soil wth complete nutrient solution exhibited a
of fructans presented here were lower than those higher shoot/root ration than -N, -P and control plants.
previously found in V. Herbacea (Carvalho & Dietrich, In his review, Wilson (1988) mentions that some
1993), possibly because, in the present investigation, authors have found no change in this ration when
younger plants were used. nutrient levels are varied, and this has usually not
been accompanied by and effect on growth, indicating
It is known that many species well adapted to
that the nutrient was not deficient in any of the
unfavorable conditions grown better when cultivated in
treatments. As a similar value of shoot:root ratio was
soil with high nutrient levels. In fact theirs ecological
found for -N, -P and control plants and growth was not
preference is due to a low competitive capacity and a
significantly affected, it can be suggested that the
high tolerance to extreme conditions (Fitter & Hay,
levels of these elements naturally found in cerrado
1983). In V. hebacea, a good supply of nutrients, as
soils are sufficient for the growth of the plant.
provided by the complete Hoagland solution seems to
promote shoot growth at the expense of fructan Considering the two substrates used in this
accumulation, a fact that most probably would treaten investigation, no significant differences in shoot length
the survival of the species in cerrado conditions, were found in plants grown in cerrado soil. However,
where accumulation is essential to endure the dry treatment with complete Hoagland solution and
season. Hoagland solution without phosphorus promoted

R. Bras. Fisiol. Veg., 9(2):89-96, 1997


... Growth and frutactan contents ... 95

shoot growth in plants cultivated in sand. The same


was observed for leaf area (Fig. 3B). REFERENCES
Archbold (1938), working with rye leaves, reported ADÁAMOLI, J.; MACEDO, J.; AZEVEDO, L.G. &
that N deficiency leads to an increase in fructan MADEIRA NETTO, J. Caracterização da região dos
contents. Firstly, levels of soluble sugars are increased cerrados. In: GOERDERT, W.J. ed. Solos dos
possibly due to the lack of nitrogen for protein cerrados: Tecnologias e estratégias de manejo. São
synthesis an, secondly, these sugars are used in the Paulo, Editora Nobel, 1985. p.33-74.
synthesis of fructans, promoting a change in the ANDREWS, P. The gel filtration behaviour of proteins
proportion of these sugars. More recently Tsukihashi related to their molecular weights over a wide range.
et al. (1991), working with yacon (Polymnia Biochemical Journal, 96:595, 1965.
sonchifolia), reported a slight decrease in total ARCHBOLD, H.K. Physiological studies in plant
oligofructans with a nitrogen supply. In the present nutrition. VII. The role of fructosans in the
study, treatment with nutrient solution without nitrogen carbohydrate metabolism of barley plant. Par t 2.
did not promote growth of rhizophores in sand or Seasonal changes in the corbohydrates with a note
growth of plants in either substrates but produced on the effect of nitrogen defficency. Annals of
small amounts of fructans when compared to control Botany, 2:403-436, 1938.
plants, specially fructo-polysaccharides. ARENS, K. As plants lenhosas dos campos cerrados
It is interesting to note that the control treatment in com flora adaptada às deficiências minerais do solo.
which nitrogen was absent (sand) or was present in In: I SIMPÓSIO SOBRE O CERRADO. São Paulo,
small amounts (cerrado soil) led to a considerable Edgar Blücher, 1963. p.285-303.
accumulation of fructans. Although these data may DELHAIZE, E. & RYAN, P.R. Aluminum toxicity and
seem contradictory, other factors may be involved in tolerance in plants. Plant Physiology, 107:315-321,
the accumulation process, like the competition among 1995.
mineral nutrients, sink organs and the possible EITEN, G. The cerrado vegetation of Brazil. Botanical
adaptive mechanisms of this species to oligotrophic Review, 38:201-341, 1972.
condition of cerrado soils. FERRI, M.G. Transpiração de plantas permanentes dos
cerrados. Boletim FFCL-USP, 41 Botânica
Phosphorus deficiency was reported to increase the
4:159-224, 1944.
soluble sugar levels and reduce vegetative growth
without affecting assimilation rate. In general, fructan FITTER, A.H. & HAY, R.K.M. Environmental
accumulation is favored and increase in the ratio physiology of plants. London, Academic Press,
fructan and total sugar is shown (Russel, 1938). these 1983. 355p.
results obtained for barley plants are different from the HOAGLAND, D.R. & ARNON, D.I. The water-culture
data presented here for V. herbacea in which method for growing plants without soil. University of
vegetative growth and fructan accumulation were not California Agricultural Experimental Station, 1938.
specially affected by the absence of phosphorus. (circular 347)
Considering the high value of shoot/root in plants GOODLAND, R. Oligotrofismo e alumínio no cerrado. In:
growing in sand and receiving complete Hoagland II SIMPÓSIO SOBRE O CERRADO, São Paulo,
solution and -P treatment and the low ration in those 1971. p.44-60.
receiving water or -N treatment, it can be suggested JERMYN, M.A. A new method for the determination of
that N is an important element affecting growth in ketohexoses in presence of aldohexoses. Nature,
these plants. 177:38-39, 1956.
KANAYA, K.I.; CHIBA, S. & SHIMOMURA, T. Thin-layer
The data here obtained indicate that nutrient
chromatography of linear oligosaccharides.
condition is one of the factors defining growth and
Agronomical Biological Chemistry, 42:1947-1948,
accumulation patterns in this species, although control
1978.
plants growing in cerrado soil and in sand presented
adaptive mechanisms that allowed efficient fructan MENEZES, N.L.; MULLER, C. & SAJO, M.G. Um novo e
peculiar tipo de sistema subterrâneo em espécies de
accumulation in the rhizophores, similar to what is
Vernonia da Serra do Cipó (Minas Gerais, Brasil).
found in plants growing in the cerrado covered
Boletim de Botânica da Universidade de São Paulo,
environment. 7:33-38, 1979.
MONASTERIO, M. & SARMIENTO, G. Phenological
ACKNOWLEDGMENTS strategies of plant species in the tropical savanna
To Drs. Rita de Cassia L. Figueiredo-Ribeiro and Gil and the semi-deciduous forest of the Venezuelan
Martins Felippe for a critical review of the manuscript. Llanos. Journal of Biogeography, 3:325-355, 1976.
POLLOCK, C.J. Fructans and the metabolism of sucrose
in vascular plants. New Phytolosist, 104:1-24, 1986.
R. Bras. Fisiol. Veg., 9(2):89-96, 1997
96

POLLOCK, C.J. Seasonal patterns of fructan and SHIOMI, N. Structure of fructopolysaccharide


metabolism in forage grasses. New Phytologist, (Asparagosin) from roots of asparagus (Asparagus
83:8-15, 1979. officinalis L.). New Phytologist, 123:263-270, 1993.
RAWITSCHER, F.K. Algumas noções sobre a TSUKINHASHI, T.; ASAMI, T.; MINASAWA, K. &
transpiração e o balanço d’água de plantas KUBOTA, M. Effect of nitrogen and potassium
brasileiras. Anais da Academia Brasileira de fertilizer on the growth and yield of yacon ( Polysmnia
Ciências, 14:7-36, 1942. sonchifolia ). Vocacional Agriculture, 38:50-56,
RUSSEL, R.S. Physiological studies in plant nutrition. I. 1991.
The effect of mineral defficiency on the fructosan WILSON, J.B. A review of evidence on the control of
metabolism of the barley plant. Annals of Botany, shoot:root ratio, in relation to models. Annals of
2:865-882, 1938. Botany, 61:433-449, 1988.
SARMIENTO, G. The ecology of neotropical savannas. WISE, C.S.; DIMLER, R.J.; DAVIS, H.A. & RIST, C.E.
Cambridge harvard University Press, 1984. 235p. Determination of easily hydrolyzable fructose units in
dextran preparation. Analytical Chemistry, 27:33-36,
1955.

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