Agriculture, Ecosystems and Environment, 13 (1985) 301--307

301

Elsevier Science Publishers B.V., Amsterdam -- Printed in The Netherlands

OBSERVATIONS ON THE VEGETATIVE GROWTH PATTERN OF

S P E A R G R A S S ( I M P E R A T A C Y L I N D R I C A (L.) B E A U V . )

A.O. AYENI

Department of Agronomy, University of lbadan, Ibadan (Nigeria)

(Accepted for publication 28 March 1985)

ABSTRACT
Ayeni, A.O., 1985. Observations on the vegetative growth pattern of speargrass (Imperata
cylindrica (L.) Beauv.). Agric. Ecosystems Environ., 13: 301--307.
The vegetative growth pattern of speargrass (Imperata cylindrica (L.) Beauv. var.
major) was studied in the greenhouse. Careful observation of plants grown from mature
rhizome fragments weighing approximately 0.4 g indicated that a young plant initiates
new rhizomes between the third and fourth leaf-stage. Rhizome growth is determinate
(the rhizome tip turns up to form a shoot), with the apical bud forming a shoot and
sub-apical buds forming rhizome branches. Under favourable conditions, apical and
sub-apical buds develop simultaneously, but under stress the growth of the apical bud
is favoured. Root and bud development occurs on the distal nodes of the young rhizome
long after the rhizome has been formed. The significance of this growth pattern in relation to speargrass as a problem weed is discussed.

INTRODUCTION
In r e c e n t y e a r s t h e role o f w e e d b;.ology in designing b e t t e r w e e d c o n t r o l
strategies has e n g a g e d t h e a t t e n t i o n o f w e e d scientists m o r e seriously t h a n
ever b e f o r e . T h i s is a p p a r e n t l y p r o m p t e d b y r e a l i z a t i o n o f t h e l i m i t a t i o n s
o f c h e m i c a l c o n t r o l m e t h o d s . E v e n t h o u g h t r e m e n d o u s success has b e e n
a c h i e v e d w i t h this a p p r o a c h , t h e r e are still s o m e p r o b l e m w e e d s w h i c h
d o n o t y i e l d to c h e m i c a l t r e a t m e n t , c a n n o t be c o n t r o l l e d a t e c o n o m i c
levels w i t h a f e w e f f e c t i v e c h e m i c a l s , o r are d e v e l o p i n g strains w h i c h are
r e s i s t a n t to c o n v e n t i o n a l r e c o m m e n d a t i o n s ( J e n s e n a n d B a n d e e n , 1 9 7 9 ) .
M a n y investigators (Sagar, 1 9 6 8 ; P a r k a , 1 9 7 6 ; H o l m et al., 1 9 7 7 ; H i l t o n ,
1 9 7 9 ) n o w share t h e v i e w t h a t a s o u n d k n o w l e d g e o f t h e b i o l o g y o f rec a l c i t r a n t w e e d s will h e l p in i d e n t i f y i n g stages in t h e i r life c y c l e at w h i c h
c o n t r o l m e a s u r e s c o u l d be d i r e c t e d in o r d e r t o a c h i e v e g o o d results.
Speargrass (Imperata cylindrica (L.) Beauv.), an i m p o r t a n t w a r m - c l i m a t e
r h i z o m a t o u s grass (Soerjani, 1 9 7 0 ; H o l m et al., 1 9 7 7 ) , b e l o n g s t o t h e g r o u p
o f w e e d s w h i c h a t p r e s e n t c a n n o t b e e c o n o m i c a l l y c o n t r o l l e d w i t h herbicides. F o r i n s t a n c e , a m i n i m u m o f o n e foliar s p r a y o f 4 kg a.i. h a -1 o f giy-

0167-8809/85/$03.30

1985 Elsevier Science Publishers B.V.

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phosate or split applications of 15--20 kg a.i. ha -1 of dalapon are required
to give effective control for a period of 1 year or less (Ivens, 1973). In
Nigeria, when the cost of application is added to the price of the herbicide,
the cost/returns analysis becomes unfavourable, especially to small-holder
farmers working less than 3 ha per annum. For weeds like speargrass, it is
thus worthwhile to pay some attention to their mode of development to
identify the "soft spots" in their life cycle. The objective of this work was to
study the speargrass growth pattern carefully in order to identify those aspects of its development t h a t need critical investigation in relation to the
problem of its control.
MATERIALS AND METHODS
This report is based on observations made between August 1976 and
June 1978 under greenhouse conditions at Muenscher Laboratory, Cornell
University, Ithaca, New York. The rhizomes from which the speargrass was
established were obtained from Auburn University, Alabama. This variety
of speargrass was identified as major (Dickens, 1974). Mature rhizome
fragments, with visible buds and weighing approximately 0.4 g, were used
for propagation in a peat--vermiculite medium supplied with adequate
fertilizer levels. Plants were raised in metal flats 48 34 X 10 cm, with
propagules planted 2--3 cm deep. Greenhouse temperatures during the
period of observations fluctuated between 21C in winter and 32C in
summer, and plants developed under natural light and day length. Observations were made at successive leaf stages beginning from the second
fully expanded leaf stage. On each occasion an average of five plants at
a similar growth stage were carefully uprooted and examined.
OBSERVATIONS AND DISCUSSION

Shoot
Speargrass (Fig. 1) is an erect rhizomatous grass. Under the conditions
of this study, the tallest plants attained a height of about 75 cm. The shoot
is a cylindrical culm formed from several leaf sheaths rolled tightly around
one another. At the centre of this " c y l i n d e r " , and usually beneath the
ground surface, is the apical meristem which is well protected from external
hazards. The leaf sheath is hairy and terminates in a green leaf blade which is
linear lanceolate. The hairs of the sheath extend only to the proximal end of
the leaf blade, and here they are restricted to the margins. Hairs are absent
towards the distal end of the leaf blade, but the margins have fine serrations.

Rhizome
Between the third and fourth leaf stage, rhizome development starts at the
base of the shoot. The number of rhizomes varies from one to four. Initially,

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Shoot

~'oung rhizome
Propagu[e~

)rous r o o t s

Fig. I. Four-leaf stage speargrass plant with a young rhizome at the plagiotropic growth
stage.

rhizome growth is plagiotropic (grows at an angle to the mother plant) (Fig.

2a). At this growth stage, the rhizome is tightly covered by whitish and
succulent scale leaves, otherwise called cataphylls (Etter, 1951). Roots are
absent on this category of rhizomes, and a close examination after removing
the cataphylls revealed that they have no visible sub-apical buds. By the fifth
leaf stage, rhizome growth becomes diageotropic (horizontal) (Fig. 2b).
A few roots are formed on the distal nodes and the cataphylls on the proximal nodes are loosely attached, darker in colour and become more fibrous.
When cataphylls are removed at this stage, sub-apical buds are visible on the
distal nodes. Between the fifth and sixth leaf stage, rhizome growth is
negatively orthogeotropic {tip turns vertically upwards) (Fig. 2c). The apical
bud subsequently forms an aerial shoot (second generation shoot), while
some of the sub-apical buds form new rhizomes (second generation rhizomes) and others, much later in the life of the second generation shoot,
develop into secondary shoots (Fig. 3a and b). Rhizomatous plants which
have rhizomes with tips turning upwards to form shoots have been described
as having determinate rhizomes (Troughton, 1957). Fisher (1965) observed
that Kentucky bluegrass (Poa pratensis L.) belongs to this group, and the
present study showed that speargrass also belongs to the same group.

304
Base of aerial shoot...
" " ~ I~
Propagule

Ground level

~ ~l~ll-~First
generation rhizome
~ " / / / / ~ ~'L (at plagiotropic stage)

Fibrous roots

Lll,

\\
(a)

Ground level
First generation
rhizome (at
diageotropic stage)

I.
(b)

Adventitious roots
(on distal nodes of
rhizome)

Ground [eve]
First generation
rhizome (at
negatively .
orthogeotrop~c
stage)

(C)

Fig. 2. Early stages of rhizome development in speargrass: (a) plagiotropic; (b) diageotropic; (c) negatively orthogeotropic growth stages. (Note that roots are absent on the
rhizome at the plagiotropic growth stage.)

Base of aerial shoot

\
ou00 .v.,

second generat,on shoot

I f
~ i /

Secondary shoot

I If

(a)
(b)
Fig. 3. Advanced stages of rhizome development in speargrass: (a) recently emerged
second generation shoot from the apical bud of a mature rhizome; (b) mature rhizome
with well-developed second~eneration rhizomes and a secondary shoot. (Note that
the second-generation rhizomes develop on the convex side and the secondary shoot
on the concave side of curvature in the mother rhizome.)

305
R h i z o m e buds

Buds are concentrated at the nodes in the apical region of the mature
rhizome. In general, bud size decreases proximally and it is estimated that
n o t more than 60% of the nodes carry visible buds. The reason for the
limited n u m b e r of buds and their restriction to the distal end of the rhizome
is n o t known, b u t observations suggest it m a y be connected with the physiological maturity of the rhizome. At the plagiotropic growth stage (Fig. 2a),
the rhizome has no sub-apical buds, b u t buds become visible from the
diageotropic stage onwards. This area of rhizome development in speargrass
deserves to be investigated. Etter (1951) observed this pattern of development in K e n t u c k y bluegrass, although it is probably n o t quite as striking
as it is in speargrass.
Root

The r o o t system of speargrass is fibrous and development occurs at the

rhizome nodes. R o o t population is high at the base of the aerial shoot
because nodes are closely packed in this region (Fig. 1). R o o t formation
on the rhizome does n o t occur until the diageotropic growth stage is reached. At the older stages of rhizome development, roots are formed on the
more distal nodes, b u t none on the proximal one. It is worthwhile investigating the cause of the delay in r o o t formation until the diageotropic stage
is reached and whether or n o t there is a connection between r o o t and
sub-apical bud development, both of which seem to start at a b o u t the same
stage of rhizome development. A knowledge of what suppresses bud development on the proximal nodes may help in understanding h o w bud developm e n t could be suppressed in the apical region o f the rhizome, which in
turn should help to substantially reduce the regenerative ability of speargrass.

Second generation shoot and rhizomes

In vigorous plants the formation of a second generation shoot and rhizome branches from a mature rhizome occur simultaneously. In weak
plants, however, the development of the aerial shoot supersedes rhizome
production. The sub-apical buds in the latter case form rhizomes only
after the second generation s h o o t is well established.
Only the buds on or towards the convex side of curvature of the mature
rhizome form new rhizomes (Fig. 3b); those on the concave side either
develop into shoots long after rhizome branches have been established
or remain permanently suppressed. The reason for this geometric specificity should be investigated. Adequate understanding of this behaviour
should facilitate the identification of how more buds could be made to
form secondary shoots and less to form rhizomes, so that the means of
propagation and persistence in speargrass is diminished.

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Tillering, in the generally accepted sense of the terminology, does n o t

occur in speargrass. Observations on var. major at Cornell University and
vat. africana under field conditions in Nigeria indicated that what might be
described as tillers (see Sajise, 1972) are actually secondary shoots developing from sub-apical buds which are present on the closely packed nodes
at the base of the primary shoot. When many such shoots develop from
the same rhizome, this results in a tuft of shoots which, w i t h o u t close
observation, could be mistaken for tillers.
CONCLUSIONS

An a t t e m p t has been made to observe speargrass vegetative growth patterns closely, with emphasis on rhizome and root development. F r o m the
study, the following questions were raised:
(a) Why does the development of buds and roots on the rhizome start long
after rhizome formation, and why are these organs (buds and roots}
absent on the proximal nodes?
(b) What causes the geometric specificity of rhizome and sub-apical (secondary) shoot development on the mother rhizome?
It is suggested that the genotypic attributes of speargrass and the m o v e m e n t
of growth substances and p h o t o s y n t h a t e s account for these peculiar characteristics. These aspects of the biology of the plant need to be carefully
investigated so that a better approach to its control can be evolved, which
will be both reliable and economical to the farmers of the humid and subhumid tropics who suffer substantial losses due to speargrass interference
with cropping activities.
ACKNOWLEDGEMENTS

The author wishes to thank Dr. Ray Dickens of Auburn University,

Alabama, for providing the propagules from which the plants used for the
experiment were raised, and the Department of Agronomy, Cornell University, for providing the research facilities.

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