Weed Research, t974. Volume 14.

365-369

Experiments with Cyperus rotundus L.

I. Growth and development and efTecls of 2,4-D and paraquat

JOHN L. HAMMERTON Facu/r>'a/ Introduction

Agriculture, University of The West Indies, Mona,
Kingston 7, Jamaica Cyperus rotundus L., purple nutgrass or nutsedge is
now recognized as the most serious weed species in
Received 4 January 1974 world agriculture (Holm & Herbergcr, 1969). Its
distribution is pantropical. extending into sub-
Summary: R^ume: Zusammenfassung tropical and other warmer regions (Holm, 1969). It
Tubers of Cyperus roturtdus L. were planted in pots, and
occurs in cultivated land and other disturbed sites,
growth and development followed by washing out every 3 and can render crop production extremely difficult.
days. New tubers were lirst recorded 16 days after shoot Apart from competitive efTects, substances toxic to
emergence began, and tuber production proceeded at ihc rate other plants may be produced by decay of dead
of one every 4 days. By 51 days a single tuber had produced
approximately nine new tubers, nine shoois and nearly 90 cm residues of C. rotutidiis (Friedman & Horowitz,
of rhizome. Treatment with 2,4-D or paraquat at 34 days 1970; Horowitz and Friedman, 1971). Hauser
severely checked shoot growth, but tuber production
appeared to he only temporarily checked. Parent luber dry (1962a, b) and Horowitz (1965, 1972) among others
weight fluctuated between 017 and 0-55 g and the parent have described its growth and development. Kasasian
tuber appears to aci as a source of food in times of stress. (1971) has reviewed much of the information on its
biology and control.
ExpMences sur le Cyperus rotundus L. /. Croissance et d^-
vehppement: effets du 2,4-D et du paraquat The work described herein was undertaken pri-
Des tuberules de Cyperus rolundus L. ont et^ plant^s dans des
marily to provide basic data to aid the development
pots d'oCi ils ont fite retires tous les 3 jours pour en suivre la of control programmes for nutgrass in Jamaica and
croissance et le dtveloppement. On a enregistr6 Tapparition other territories of the Commonwealth Caribbean.
de nouveaux tubercules 16 jours apris le dtbut de la levte. La
production de tubercules s'est poursuivie ^ raison d'un tous The effects of two common herbicides on nutgrass
ies 4 jours. Au bout de 51 jours, un tubercule avait produit development were also studied.
approximativement 9 nouveaux tubercules, 9 pousses et pres
de 90 cm de rhizome. Le trailement avec le 2,4-D et le para-
quat au bout de 34 jours a provoqu^ un freinage important de
la croissance de la partie a^rienne mais la production des Experimental methods
tubercules n'a ct6 ralentie que temporairement. Les poids sees
des tubercules initiaux ont varid entreO,l7 get 0,55 get ces
tiihercules paraisscnt jouer le rdle d'une source de maliircs Nutgrass was grown in 16-cm pots holding approxi-
nutritives pendant les p6riodes difticiles. mately 19 1 of sieved, methyl bromide-fumigated
loam (Maverly series). Nutgrass tubers freshly dug
Versuche mit Cyperus rotundus L. I. Wachslum und Entwick- from one site at Mona were used. Tubers were
lung und Wirkung von 2,4-D und Paraquat
sorted and trimmed of roots, rhizomes and shoots:
Knollen von Cyperus rotundus L. wurden in GefSsse gepfianzt
und das Wachstum und Enlwicklung durch Auswaschen im only sound undamaged tubers between 8 and 12 mm
Abstand von jeweils 3 Tagen verfolgt. Neue Knollen wurden in size (greatest dimension) were used. Four tubers
zuerst 16 Tage nach dem Beginn der Sprossbildung beobachtet. were planted 2-5 cm deep in each pot. Germination
Alle 4 Tage wurde eine weitere Knolle gebildet. Nach 51
Tagen hatte eine einzelne Knolte etwa 9 neue Knollen und was rarely less than 100"<, and pots with misses 8
fast 90 cm Rhiirom produziert. Eine Behandlung mit 2.4-D days after planting were discarded. Pots were kept
Oder Paraquat nach 34 Tagen hemmte das Sprosswachstum outside in full sunlight and watered as necessary to
stark, atier die Knollenbildung schien nur vorubergehend
gehemmt zu sein. Das Trockengewicht der Mutterknollen keep the soil moist.
schwankte zwischen 0,17 und 0,55 g. Die Mutterknolle The experiment was planted on 20 August 1970.
scheint in Zcilen in denen die Pflanzc Belaslungen ausgesetzt
ist als NahrstolTquelle zu wirken. At 3-day intervals up to 51 days three pots selected
 366 John L. Hammerton

at random were washed out under a jet of water to

remove soil from the plant system, which was then
separated into component parts. On 23 September
(34 days) two sets of eighteen pots were removed;
one was sprayed with paraquat dichloride at the
equivalent of 0-28 kg/ha (ion), and the other with
2,4-D (dimethylamine salt) at 112 kg/ha (a.e.).
At intervals, three randomly chosen pots from each
set were washed out and the plant system separated.
The plant system was separated into shoots, basal
bulbs (leaf bases or corms), parent (i.e. planted)
tubers, 'new' tubers (both those with shoots and
those without), rhizomes and roots. Shoots, tubers,
rhizome segments and inflorescences were counted,
and total rhizome length recorded. Dry weights were
obtained by oven drying at 90°C. Results are
expressed on a 'per parent tuber' basis.

Results

Data from the 'untreated' series are summarized in

Fig. 1. Shoots appeared above ground on day 5
(where day 0 is the day of planting) and increased to
about nine per tuber over the 51-day period. Most
of the later shoots were produced by horizontal
rhizomes from basal bulbs of the original or older Fig. I Changes with time in (a) number of shoots (•) and
inflorescences ( x ) ; (h) number of new tubers; {c) dry weight
shoots, but some arose still later from new tubers. (g) of shoots {•) and new tubers ( x ) ; (dl dry weight (g) of
The relationship of shoot number {s) to time in days roots (•) and rhi?omes(x ); (e) dry weighl(g) or basal bulbs;
(f) dry weight (g) of parent tubers; (g) toial rhizome length
((/) is given by .r = 1O+OO5W+OOO2£/^ Shoot dry (cm). All data are expressed 'per parent (i.e. planted) tuber'.
weight per planted tutwr increased linearly, the Vertical lines in (f) are fiducial limits (P = 005).
equation being i« = 0 O67f/-O-59. Inflorescence
initials were visible at 21 days and by 24 days scapes roots and horizontal rhizomes. Basal bulbs increased
had elongated. Inflorescence numtier increased curvilinearly in dry weight, the equation being
linearly to about day 39, then remained more or less k = 0 1 7 - 0 O15£y+OOOI4f/^ The number of basal
constant. New tubers (>5 mm) were present at day bulbs paralleled that of shoot number. Parent
21 (16 days after shoots first appeared) and tuber tubers showed no consistent trend in dry weight:
number increased linearly until 51 days (? = O-23d- there was no immediate drop in dry weight following
3-8) when about nine per planted tuber were present. germination but a minimum of 0-17 g per tuber was
The dry weight of new tubers also increased linearly reached at 15 days. Subsequently an increase in dry
to just over 1 g per planted tuber. The equation is weight occurred, although this was checked at
r, =OO37/-O-73. Total rhizome length (r,) in- 36-45 days.
creased at the rate of I -78 cm per day per viable The tips of shallow (2-3 cm deep), horizontally
planted tuber (r, = ]-78rf-9-4), and rhizome dry growing rhizomes in most cases turned upwards to
weight also showed a linear increase C^, = 0QO62d- produce a leafy shoot, simultaneously producing a
0056). small swollen basal bulb from which adventitious
Root production by planted tubers had started at roots soon grew. In other instances this swelling
3 days; root dry weight increased linearly to about was much larger, hard and dark in colour, resembling
0-7 g per tuber (y^ = OOiSd = 0 05). By day 6 the a tuber from which arose, apparently directly, a
vertical rhizome from viable planted tubers was leafy shoot. The tuberous swellings were usually
swelling just below the soil surface to form a basal elongated horizontally and up to 12 mm maximum
bulb, which by day 9 was producing adventitious dimension. No anatomical studies were made, but it
 Experiments with Cyperus rotundus L. 367

Table 1 Viability of various underground organs of C. are shown in Fig. 2. By day 40 (6 days after spraying)
rotundua when planted in moist vermiculitc. (The duration of 2.4-D-treated plants were chlorotic and shoots and
Ihe test was 21 days.)
scapes abscissed readily. This abscission was also
true of subterranean leafy shoots suggesting trans-
Number Number X location. The dry weight of wholly or predominantly
planted giving a viability
shoot green leafy shoots decreased with time. Tbis was due
largely to necrosis of older shoots as new shoots
were small and many died soon after emergence.
Basal bulbs 60 48 80 Old basal bulbs failed to tiller. Thirty-one days after
'Tuberized' basal bulbs* 60 35 58 spraying shoot number was only half that at spray-
New dormant black tubers 30 10 33
New dormant white tubersf 40 12 30 ing.
Deep tubers with shootsj: 40 22 55 Tuber number showed a decrease after spraying:
Old parent tubers 40 30 75 many tubers initiated shoots which abscissed or
became necrotic and most of these tubers decayed.
* See icxt for description. Tuber number and weight, however, increased from
t Young immature tubers. 45 days in spite of a considerable loss of green
j Tubers apparently producing a leafy shoot directly which
failed to reach the soil surface. pbotosynthetic tissue. Some tubers showed a con-
striction suggesting a check to growth. Old basal
bulbs showed little cbange in dry weight and did not
seems likely that these structures were fused tubers decay but produced no new shoots. The weight of
and true basal bulbs. Their occurrence was not 'new' basal bulbs remained unchanged with time
apparently related to depth. They have been referred
to as 'lubcrized' basal bulbs and in part account for
the increase in basal bulbs dry weight from 36 days.
Deeper rhizomes produced tubers and tuber chains.
Chains with more than four tubers were not found.
Tubers occasionally produce two or even three
rhizomes which in turn formed tubers. A few deep
tubers gave rise, apparently directly, to leafy shoots,
thereby resembling 'tuberized' basal bulbs, without
producing a vertical rhizome. These shoots in-
variably decayed and never reached the surface.
This phenomenon could be a result of pot conditions
as. by about 50 days, some thirty-five shoots, thirty-
five tubers and 3-6 m of rhizome plus a mass of roots
were present in each pot.
Senescence of the outer leaves of the older shoots
occurred from about 36 days. 'Tillering' (production
of a new shoot apparently directly from the basal
bulb) was noted occasionally. At the conclusion of
this study various organs were placed in moist
vermiculite to assess viability. Results are summarized
in Table 1. Basal bulbs gave 80% viability within
the 21-day test period. Old parent tubers gave a
similar value but other organs showed varying Fig. 2 Effects of 2,4-D (M2 kg/ha a.c.) and paraquat (0 28
kg/ha ion) on day 34 (arrow) on (a) dry weight (g) of shoots;
degrees of dormancy. No signs of decay were present (b)dry weight (g)of new tubers; (c) dry weight (g) of rhizome;
at the end of tbe test in any organ. Both black and (li) dry weight <g) of roots; (e)dry weight (g) of parent tubers;
white new tubers (these had produced no shoots and and (0 number of new tubers. All data are expressed 'per
parent (i.e. planted) tuber'. Solid circles (•) represent means
were therefore assumed to be dormant) showed of the untreated series, with line of best fit shown; triangles
evidence of dormancy even when detached and ( . ) indicate means from the 2,4-D-treated series, open circles
planted in vermiculite. (_) means from the paraquat-treated series; interrupted lines,
for 2,4-D- and paraquat-treated series Httcd by eye. Vertical
Data from the 2,4-D- and paraquat-treated series lines in (e) are liducial limits (i* -^ 0 05).
 368 John L. Hammerton

but many shoots from these became necrotic. if tuber production is to be prevented. The data
Root proliferation was noted on tubers and basal suggest that no more than 14 days should elapse
bulbs at 43 days but is not apparent from the root between shoot emergence and cultivation or chemical
weight data. Rhizome dry weights show no clear treatment. Delay until flowering scapes and more
trend but some decay followed by recovery may have shoots are present would lead to many more tubers
occurred. Parent tuber dry weight decreased relative being produced.
to the untreated series but parent tubers did not Tbe total dry weight per parent tuber at 51 days
produce new shoots. was 715 g of which approximately half consisted of
Paraquat caused rapid necrosis of green shoots but reproductive structures (tubers and basal bulbs).
tbere was a rapid increase in shoot number, mainly This represents a growth rate of about 015 g per
small shoots arising as 'tillers' from old basal bulbs, day. It is intended to make a more detailed analysis
so that shoot number did not decline to less than two of growth in this and otber experiments later. Black,
per parent tuber. Sboot dry weights did decline, Chen &. Brown (1969) have reported C. rotu/ulus to
however, (Fig. 2a). Tuber number fell as did tuber have high photosynthetic efficiency by way of the
dry weight following treatment. Old basal bulbs C4 dicarboxylic acid pathway of COi fixation.
showed a decrease in dry weight associated with It is clear that basal bulbs, also called corms by
decay but some tillered to give small new shoots. Wills & Briscoe (1970), can act as propagules as they
Most new shoots came from young tubers. By 70 contain starch and can initiate rhizomes and shoots.
days some inflorescences were present, though shoot The so-called tuberized basal bulbs may be fused
weight was less than at 21 days. Some decay and basal bulbs and tubers: they contain presumably
loss of dry weight of rhizomes and roots occurred, greater starch reserves than ordinary basal bulbs and
and parent tubers also showed a loss of weight so would be more effective propagules under adverse
followed by recovery. conditions. The production of subterranean shoots
unable to reach the surface is not an artefact of pot
culture, as it has been observed in the field at Mona.
Discussion It is disadvantageous to the plant and no explana-
tion of its cause or function can be given.
The initial planting density of four tubers per pot is The fluctuations in parent tuber dry weight
approximately equivalent to 200 per m^ This is low suggest that they can act as sinks and sources of
compared with field infestations of 1700 to 3600 per carbohydrates, the latter presumably in times of
m^ (Hammerton, 1968). 2080 to 3200 per m' stress. Tbis is surprising as new tubers were being
(Tripathi, 1969), 1940 per m^ (Baker, 1964), 825 per produced at times of increa.se in parent tuber dry
m^ (Rao, 1968) and 1090 per m^ (Hauser, 1962b) weight. Possibly the pots constrained tuberization,
reported from elsewhere. Investigations at Mona so leading to an excess of photosynthate. Field
have given values up to 8700 tubers per m^ (Ham- studies (Hammerton, unpublished data) have also
merton, unpublished data). The final tuber density shown large fluctuations in mean tuber dry weight,
of 2000 per m* is not high compared with many of although no separation into old and new tubers was
these values. made. Although parent tubers bad remained in-
Of major importance is tbe earliness and rapidity active for some 48 days after having produced the
of tuber production. New tubers were present only first shoot, only 25% showed any dormancy when
16 days after shoots appeared above ground: at this tested.
time shoots had only six leaves. From then on until The effects of 2,4-D on nutgrass have been reported
the experiment finished a new tuber was produced by Thakur (1952). Muzik & Cruzado (1953). Eames
every 4 days approximately. Rao (1968), however, (1949), Sinha & Thakur (1966) and others. Abscis-
reported four new tubers within 30 days and ninety- sion and root proliferation are well-known effects of
nine in 90 days. Hauser's (1962a) studies showed that phenoxy herbicides. The effects on shoot number and
tuber formation did not begin until 6-8 weeks after dry weight and on basal bulb dry weight in the
initial foliar emergence. In the present experiment present work are striking. The impact on tuber
sboot and tuber production proceeded concurrently production is comparatively small, however: a delay
throughout the period. in tuber production of 10 days or so appeared to be
It is clear that control measures must be under- followed by resumption of tuber production at a
taken extremely early and at relatively short intervals rate similar to the original. A somewhat similar
 Experiments with Cyperus rotundus L, 369

sequence followed treatment with paraquat. The EAMES A.J. (1949) Comparative effects of spray treatments
with growth regulating substances on nutgrass. Cyperus
drop in parent tuber dry weight relative to the un- rotundus L., and anatomical modifications following
treated series suggests that parent tubers supplied treatment with butyl 2,4-dichlorophenoxyacctate. Am. J.
much of the energy for shoot and new tuber pro- Bor.. 36, 571-584.
FRIEDMAN T . & HOKOWITZ M . (1970) Phytotoxicity of sub-
duction. Muzik & Cruzado (1953) reported very terranean residues of three perennial weeds. Weed Res., 10,
slow movement of 2,4-D in nutgrass systems result- 382-385.
HAMMERTON J.L. (1968) Nutgrass in Panama: first impression.
ing in a limited kill. PANS (C). 14, 339-345.
Paraquat rapidly destroyed shoots and many HAUSER E . W . (1962a) Establishment of nutsedge from space-
basal bulbs. Regeneration was rapid, however, planted tubers. Weeds, 10, 209-212.
HAUSER E.W. (1962b) Development of purple nulsedgc
compared with the protracted efFect of 2,4-D, under field conditions. Weeds, 10, 315-321.
although much of the new shoot growth came from HOLM L . G . (1969) Weed problems in developing countries.
new basal bulbs or new tubers. The effects on root Weed Sci., n, 113-118.
HOLM L.G. & HERBERGCR. J. (1969) The world's worst weeds.
and rhizome dry weight are surprising in view of the Proc. 2nd Asian Pacific Weed Control Interchange, 1-14.
poor translocation of paraquat in bright sunlight HoRowiTZ M. (1965) Data on tbe biology and chemical
(Ashton & Crafts, 1973). Parent tubers again control ofthe nutsedge (Cvperus roiundus) in Israel. PANS
(C), 11. 389-416.
appeared to function as a source of energy for new HOROWITZ M . (1972) Growth, tuber formation and spread of
growth. Cyperus rotundus L. from single tubers. Weed Res., 12,
348-363.
In conclusion, this work suggests an important HOROWITZ M . & FRIEDMAN T . (1971) Biological activity of
role for the parent tuber apart from the fact that it subterranean residues of Cynodon daciylon L., Sorghum
can produce further shoots from dormant buds if the halepense h. and Cyperus rotundtts L. Weed Res., 11,88-93.
KASASIAN L. (1971) Weed Control in the Tropics, pp. 83-90.
current shoot is destroyed. This merits further work Leonard Hill, London.
but may explain why certain herbicides, such as MUZIK T . J . & CRUZADO H.J. (1953) The elTcct of 2,4-D on
glyphosate and possibly organo-arsenicals, are more sprout formation in Cyperus rotundus. Am. J. Bot., 40,
507-512.
effective in nutgrass control than 2,4-D in that they RAO J . S . (1968) Studies on the development of tubers in
are translocated further. nutgrass and their starch content at dilTerent depths of soil.
Madras agric. J., 55, 19-23.
SiNHA T.D. & THAKUR C . (1966) Eradication of nutgrass by
References killing Ibe tubers with 2,4-D. Madras agric. J., 53, 72-75.
THAKUR C. (1952) Effect of MCPA on nutgrass. Agron. 7., 44,
ASHTON F.M. & CRAFTS A . S . (1973) Mode of Action of 589-590.
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BAKER R.S. (1964) Reproductive capacity of nutsedge Population of tubers at dilferent depths of the soil and their
(Cyperus rotwidusHnbers. Abstr. Meeting Weed Soc. Amer., sprouting response to air drying. Proc. Nat. Acad. Sci.
63-64. India (Seel. B). 39, 140-142.
BLACK C . C , CHEN T . M . & BROWN R . H . (1969) Biochemical WILLS G . D . & BRISCOE G . A . (1970) Anatomy of purple
basis for plant competition. Weed Sci., 17, 338-344. nutsedge. Weed Sci., 19, 63\-6iS.