Published online October 3, 2006
Allelopathic, Autotoxic, and Hormetic Effects of Postharvest Sugarcane Residue
Ryan P. Viator,* Richard M. Johnson, Casey C. Grimm, and Edward P. Richard, Jr.
ABSTRACT
With green sugarcane (interspecific hybrids of Saccharum spp.)
harvesting, 6 to 24 Mg ha21 of postharvest residue is deposited on the
field surface covering the sugarcane stubble that must reemerge for
several ratoon crops. The objectives of this research were to: (i)
Reproduced from Agronomy Journal. Published by American Society of Agronomy. All copyrights reserved.
determine if postharvest residue possesses allelopathic, autotoxic, and
hormetic properties; (ii) determine the interaction of soil type with
possible autotoxic effects; and (iii) identify a reliable indicator species.
Extract concentrations consisted of 0, 0.1, 10, 25, and 100% of the
original solution of a 1:28 tissue to water extract. The higher con-
centrations of residue extracts exhibited autotoxicity by delaying early
leaf development. The lower extract concentration of 10% increased
sugarcane bud germination by 45% compared with the control, in-
dicating hormetic effects. Allelopathic activity on tall morninglory
(Ipomoea hederacea Jacq.) was more pronounced on a light soil;
germination and radical length were reduced by all concentrations by
an average of 42% and 8 mm, respectively, compared with the control.
Seedling dry weights were reduced by an average of 10 mg by the
10, 25, and 100% extract concentrations relative to the control. On
the heavy soil, only the 100% concentration reduced radical length
and weight by 5 mm and 4 mg, respectively, relative to the control.
Extract effects on oat (Avena nuda L.), rye (Secale cereale L.), and
tomato (Lycopersicon esculentum Mill.) showed poor correlation
with effects on sugarcane. Chemical analysis by gas chromatography/
mass spectrotometry indicated the extract contained benzoic acid.
Further studies are needed to establish the impact of benzoic acid in
natural settings.
S UGAR INDUSTRIES throughout the world are rapidly
adopting green-cane harvesting mainly due to public
pressure to reduce smoke and soot resulting from the
burning of standing sugarcane, the potential for in-
creased sugar recovery with this system, and the desire
to prevent soil degradation, erosion, and environmental
pollution (Richard et al., 2001; Sampietro and Vattuone,
2006a). With green-cane harvesting, 6 to 24 Mg ha21 of
postharvest residue, regardless of whether the sugarcane
is manually or mechanically harvested, is deposited on
the field surface. Postharvest residues consist of non-
mature stalk materials (such as green leaves and im-
mature internodes), senesced leaves, and small pieces
of mature stalks that result from the harvesting pro-
cess. Sugarcane must then reemerge through this debris
to produce ratoon crops, as sugarcane is a perennial
crop that is grown for 3 to 8 yr from a single planting.
R.P. Viator, R.M. Johnson, and E.P. Richard, Jr., USDA-ARS South-
ern Regional Res. Cent., Sugarcane Res. Lab., 5883 USDA Rd.,
Houma, LA 70360; and C.C. Grimm, USDA-ARS Southern Regional
Res. Cent., Food Processing and Sensory Quality Unit, 1100 Robert
E. Lee Blvd., New Orleans, LA, 70124. Received 31 Jan. 2006.
*Corresponding author (rviator@srrc.ars.usda.gov).
Published in Agron. J. 98:1526–1531 (2006).
Sugarcane
doi:10.2134/agronj2006.0030
ª American Society of Agronomy
677 S. Segoe Rd., Madison, WI 53711 USA
1526
Considerable research has been conducted concern-
ing sugarcane allelopathy. Allelochemicals have been
identified and isolated from sugarcane leaves (Singh
et al., 2003). These phytotoxic compounds include 2,4-
dihydroxi-1,4-benzoxazin-3-one and 2-benzoxazolinone.
Concentrations of these two compounds ranging from
0.45 mM to 1.25 mM reduced root growth of lentil (Lens
culinaris Medik.) seedlings, but seed germination was
not affected at these concentrations. Earlier research
suggested that phenolics may be involved in the phyto-
toxicity caused by sugarcane straw (Wang et al., 1967).
Ferulic, vanillic, and syringic acids have also been iden-
tified as phytotoxins in sugarcane straw (Sampietro
et al., 2005; Sampietro and Vattuone, 2006b). These com-
pounds increased root cell leakage, inhibited dehy-
drogenase activity, and reduced chlorophyll content in
lettuce (Sampietro et al., 2005). In another study, extracts
from green sugarcane leaves reduced the germination of
lettuce (Lactuca sativa L.) seeds, but senesced leaves
had no effect on germination of lettuce (de Carvalho
et al., 1996). Leachate from sugarcane straw that was
air-dried for 48 h at 608C contained water-soluble phe-
nolics that interfered with seedling growth of beg-
garticks (Bidens subalternans L.) and wild mustard
(Brassica campestris L.). Leachate incorporated in biotic
soil inhibited weed root growth more than leachate in-
corporated in abiotic soil, suggesting that microbial
activity is involved in allelopathic activity. Soil char-
acteristics evaluated in soil treated with sugarcane straw
leachate suggested that straw phytotoxicity is not related
with variations in inorganic nutrients (Sampietro and
Vattuone, 2006a).
In addition to allelopathic effects, autotoxicity has
been reported in sugarcane. Autotoxicity is an inter-
specific form of allelopathy that occurs when a plant
releases chemicals harmful to its growth and devel-
opment (Miller, 1996). Sterilized leachates from soils
where sugarcane was previously grown inhibited the
growth of sugarcane test plants (Magarey and Bull, 1994).
In both field and laboratory experiments, Wang et al.
(1984) found that decomposing residues contained fe-
rulic, p-hydroxybenzoic, vanillic, p-coumaric, and syringic
acid, which caused sugarcane phytotoxicity.
Allelopathy and autotoxicity are influenced by many
environmental factors. Leaching of allelopathic com-
pounds from sorghum residue in field experiments de-
pended on precipitation (Roth et al., 2000). Laboratory
studies showed that severity of alfalfa autotoxicity is
influenced by soil type and rainfall patterns (Jennings
and Nelson, 1998). Allelochemicals may be transported
through the soil and can be transformed, metabolized,
or become bound to organic matter during this process
(Inderjit, 2001a). Inconsistent allelopathic effects sug-
Abbreviations: GC, gas chromatography; MS, mass spectrotometry.
 VIATOR ET AL.: SUGARCANE RESIDUE ALLELOPATHY
gest that the severity and duration of field autotoxicity
may vary with environment and geographic location
(Jennings and Nelson, 2002). Thus, in laboratory in-
vestigations, it is critical that concentrations of chemicals
tested reflect those that exist in the soil–plant system un-
der investigation (Al Hamdi et al., 2001; Inderjit, 2001a).
Therefore, tissue to water ratios should be based on field
residue densities and average rainfall for the areas under
investigation. Moreover, allelopathy research on sug-
Reproduced from Agronomy Journal. Published by American Society of Agronomy. All copyrights reserved.
arcane is often conducted using oven-dried tissue to
make leachates (Sampietro et al., 2005; Sampietro and
Vattuone, 2006a; Sampietro and Vattuone, 2006b) even
though in field conditions nondried residue is causing
yield decline (Viator et al., 2005).
A common approach used in autotoxicity research is
the identification and use of an indicator or test species.
The indicator species and the species under investiga-
tion must show a similar response to the allelochemicals;
the indicator species allows for the advantages of uni-
form seed quality, high sensitivity, and ease of use. For
example, several researchers reported a strong correla-
tion between the allelopathic activity of rice (Oryza
sativa L.) on lettuce and ducksalad [Heteranthera limosa
(Sw.) Willd.] (Ebana et al., 2001). A reliable indicator
species would greatly enhance sugarcane bioassays be-
cause storage of sugarcane stalks for long periods leads
to deterioration, which reduces germination (Weekes,
2004) and leads to desiccation of buds (Ellis and Merry,
2004). Moreover, bud germination varies along the
sugarcane stalk after exposure to cold weather.
Preliminary field experiments in Louisiana indicated
that the yield effects from postharvest residue are in-
consistent; in some cases, residue retention did not
reduce yields. This inconsistent effect may be due to
autotoxic and/or hormetic properties of the residue
itself. Hormesis is a form of allelopathy where subin-
hibitory levels of a phytotoxic substance have stimula-
tory effects on an organism (Southman and Ehrlich,
1943; Thiamann, 1956). The objectives of this research
were to: (i) determine if fresh postharvest residue from a
sugarcane cultivar commonly grown in over 10 different
countries, ‘LCP 85-384’, possesses allelopathic, autotoxic,
and hormetic properties at residue densities represen-
tative of the Louisiana growing region; (ii) determine the
interaction of soil type with possible autotoxic and al-
lelopathic effects; and (iii) identify a reliable indicator
species for use in postharvest residue bioassays.
MATERIALS AND METHODS
Fresh sugarcane postharvest residue was collected from
four fields of LCP 85-384 (Milligan et al., 1994) on the same
day the sugarcane was harvested using a chopper harvester
(3 Dec. 2004, 20 Dec. 2004, 15 Dec. 2005, 15 Jan. 2006). LCP
85-384 was planted to 91% of the sugarcane acreage in Lou-
isiana in 2004 (Legendre and Gravois, 2004). Nine 1-m2 sections
of residue were randomly collected manually from each field
location. This postharvest residue consisted of all material that
the harvester removed from the cane stalks and deposited on
the field surface. Residue was immediately stored in burlap
sacks at 458C. On the day following each collection, a cold-
water procedure was used to extract possible allelochemicals
1527
using distilled water in a temperature-controlled water bath at
258C with a 1:28 tissue to water weight ratio (Harper and
Lynch, 1982). Temperature and residue concentrations were
based on historical weather records during the months of
residue decomposition in Louisiana (October–March). The
extract was then filtered with cheese-cloth and diluted (Ber-
geron et al., 2002). Final concentrations were 0, 0.1, 10, 25, and
100% of the original extract solution.
The aqueous sugarcane extract, consisting of 10-mL aliquots,
was washed with 2 mL of ethyl acetate, and the aqueous por-
tions discarded. A 1-mL injection of the organic phase was
made into the injector of an HP5973 gas chromatography (GC)
mass spectrotometry (MS) held at a temperature of 2708C.
Helium was employed as the carrier gas at a rate of 1 mL/min.
The GC oven temperature was initially held for 1 min at 808C
and then increased at 58C min21 to 1258C. The temperature was
further increased at a rate of 108C min21 to 3258C where the
oven was held for 4 min for a total run time of 34 min. The GC
employed a 30-m by 0.25-mm diam., with a 0.5-mm DB-5 film,
capillary column. The MS was operated in scan mode and em-
ployed electron ionization. Initial identification was made us-
ing the seventh edition of the Wiley mass spectral library
(McLafferty, 2005). Solid phase microextraction was also per-
formed directly on the aqueous extract. Aliquots of 5 mL were
placed in 10-mL vials and sealed. Samples were heated to 658C,
and a Carboxen/DVB/PDMS SPME fiber was then introduced
into the headspace of the vial. Following a 15-min adsorption
period, SPME fibers were then desorbed in the injection port
for 1 min with GC/MS run parameters as above.
To determine possible autotoxic and hormetic properties of
the extract, soil bioassays were conducted using both light- and
heavy-textured soils. Bulk samples from the plow layer (top
15 cm) of a Commerce silt loam (fine-silty, mixed, nonacid, ther-
mic Aeric Fluvaquents) soil and a Sharkey clay (very-fine, mont-
morillonitic, nonacid, thermic Vertic Haplaquepts) soil were
collected on 15 July 2003 in Lafourche Parish, Louisiana from
two fields of each soil type. Twenty samples consisting of 0.3 m2
were collected randomly from each field using a large probe.
Soils were collected from fallow ground that had been disked at
frequent intervals beginning in March to ensure destruction of
the previous ratoon crop to reduce potential accumulation of
autotoxic chemicals that may occur in soils previously cropped
with sugarcane. These fields did not have any residual herbicides
applied to them in the 14 mo before sampling. Soils from the
different fields were air-dried and sieved through a 6-mm screen
and then stored separately in sealed metal storage containers at
208C. Soils were analyzed for chemical composition by A&L
Laboratories (Memphis, TN) (Table 1). Phosphorus and major
Table 1. Chemical properties for the two soils used in the green-
house experiments investigating the interaction of soil type
with allelochemicals.
Soil classification Commerce silt loam Sharkey clay
pH† 7.1 8.0
P, mg/kg 155 34
K, mg/kg 286 101
Ca, mg/kg 4290 4063
Mg, mg/kg 759 249
S, mg/kg 18 16
Cation exchange capacity, 23.5 18.2
mol (1)/kg
Organic matter, g/kg 0.35 0.10
† Phosphorus and major cations were estimated using the Mehlich 3 ex-
traction procedure and inductively coupled plasma-atomic emission spec-
trophotometry (EPA Method 200.7), respectively. Soil organic matter
was determined by Walkley–Black oxidation (Nelson and Sommers,
1996). Soil pH was determined in a 1:1 soil to water suspension and soil
buffer pH using the SMP buffer (Thomas, 1996). The soil cation exchange
capacity was calculated by summing exchangeable cations.
 1528 AGRONOMY JOURNAL, VOL. 98, NOVEMBER–DECEMBER 2006
cations present in soil samples were estimated using the Mehlich
3 extraction procedure and inductively coupled plasma-atomic
emission spectrophotometry (USEPA Method 200.7), respec-
tively. Soil organic matter was determined by Walkley–Black
oxidation (Nelson and Sommers, 1996). Soil pH was determined
in a 1:1 soil to water suspension and soil buffer pH using the SMP
buffer (Thomas, 1996). The soil cation exchange capacity was
calculated by summing exchangeable cations.
Pots, 15 cm in diameter, were filled with 2 kg of soil. Eye
pieces of sugarcane cultivar LCP 85-384 were germinated in
Reproduced from Agronomy Journal. Published by American Society of Agronomy. All copyrights reserved.
distilled water-moistened tissue paper and then planted in pots
on 15 Jan. 2005 and 30 Jan. 2005. Pots were hand-watered daily
at a rate of 150 mL of extracts (made from the residue collected
on 3 Dec. 2004 and 20 Dec. 2004), which maintained soils near
field capacity. Leaf number and plant height were recorded
every 2 wk for 8 wk. At the termination of the experiment, final
height, leaf number, fresh weight, and dry weight were re-
corded. The experimental design was a randomized complete
block with four replications for all treatments; the experiment
was conducted twice using different residue and soil samples for
the two trials. Replication was based on the distance from an
external greenhouse wall to block for a slight temperature
gradient. Plants were grown for 8 wk in a greenhouse under
natural light with controlled relative humidity of 60 to 80% and
temperature of 30/258C (day/night).
Allelopathic properties of the extracts were determined
using tall morningglory (Ipomoea hederacea Jacq.) that was
scarified for 15 s. Similar to methods described in Sampietro
and Vattuone (2006b), 15 g of either the Commerce silt loam
or Sharkey clay soil was used as the extract absorber in 9.5-cm
sterile Petri dishes. Ten seeds were placed approximately 2 mm
into this soil layer and were then moistened with 10 mL of the
various extract concentrations (made from the residue col-
lected on 15 Dec. 2005 and 15 Jan. 2006). Dishes were im-
mediately sealed with parafilm and incubated in the dark at a
constant 268C in an environmentally controlled incubator.
Four replications/dishes were used, and the experiment was
conducted twice. Percentage germination was recorded 4 d
after incubation with radicals protruding at least 1 mm through
the seed coat considered germinated. Radical length and seed-
ling dry weight were recorded at this time.
To identify a possible indicator species, the inhibitory ac-
tivity of the various concentrations of the extract were de-
termined by seed germination and radical growth bioassays on
three test species: oat (Avena nuda L.) cultivar Rodeo, com-
mon rye (Secale cereale L.), and tomato (Lycopersicon escu-
lentum Mill.) cultivar Celebrity and sugarcane cultivar LCP
85-384. Radical length for sugarcane was not measured be-
cause radicals from the bud do not appear until several weeks
after germination; the newly germinated bud depends entirely
on roots formed from the mother stalk piece until it forms its
radicals (James, 2004). High quality seed were used; oat, rye,
and tomato each had at least 80% germination. Fifty seeds of
oat, rye, and tomato, and 10 nodal buds of sugarcane were
germinated in 9.5-cm Petri dishes on Whatman no. 541 filter
paper, with 5 mL of the various extract concentrations (made
from the residue collected on 3 Dec. 2004 and 20 Dec. 2004)
used to moisten the filter paper (Ahn and Chung, 2000; Ebana
et al., 2001). Dishes were immediately sealed with parafilm
and incubated in the dark at a constant 268C in an environ-
mentally controlled incubator. Three replications/dishes were
used for each plant species, and the experiment was conducted
twice. Percentage germination was recorded 7 d after in-
cubation with radicals protruding at least 1 mm through the
seed coat considered germinated. Radical length was also
recorded at this time using the same seedlings for oat, rye, and
tomato. Percentage sugarcane germination was recorded 14 d
after incubation with buds protruding at least 1 cm from the
stalk tissue considered germinated.
All data were analyzed using SAS with PROC MIXED (SAS
Inst., 2001) with extract concentrations and soil type as fixed
variables and trial and replication as random variables. Per-
centage data was transformed by the arc sine square root trans-
formation. Differences between treatment least square means
were compared using the pdiff option (Saxton, 1998) at the 0.05
probability level. Correlation analysis was made between the
extract effects on sugarcane and the possible indicator species of
rye, oat, and tomato. Data were also analyzed taking into
account possible hormetic effects using methods described by
Schabenberger et al. (1999) using the following equation:
a2g
E[Y|x] 5 g 1
1 1 v exp[bln(x/RD50)]
where E[Y|x] represents the average response at x dosage, a
and g are the upper and lower asymptotes of the response, v is
the initial slope, b is the point of inflection of the curve, and
RD50 is the effective dosage at which 50% of the total effect
is demonstrated.
A pseudo-r 2 value was calculated using methods described
by Schabenberger et al. (1999) with the following equation:
Pseudo-r 2 5 1 2 SSRes/SStotal(corrected)
RESULTS AND DISCUSSION
Compounds identified by GC/MS from sugarcane
extracts included: benzoic acid, decane, diacetyl glycol,
methyl hexadecanoic acid, and phthalic acid ester. Of
these compounds, benzoic acid and its derivatives have
been shown to have allelopathic properties on several
species including cotton (Gossypium hirsutum L.),
wheat (Triticum aestivum L.), ryegrass (Lolium spp.), cu-
cumber (Cucumis sativa L.), and radish (Raphanus sati-
vus L.) (Inderjit and Bhowmik, 2004; Lodhi et al., 1987;
Wu et al., 2002). Previous sugarcane research indicated
that phenolics were involved in the phytotoxicity caused
by sugarcane straw (Wang et al., 1967). Derivatives of
benzoic acids can alter leaf stomatal conductance and
transpiration, ion uptake, and net assimilation rate as
well as induce oxidative cell damage (Yu et al., 2003;
Politycka, 1996).
Generally, sugarcane development beyond germina-
tion was not effected by the extract, except for leaf de-
velopment (Table 2). Leaf number was reduced by 0.5
leaves at 2 wk after treatment by the 1.0 and 100%
concentrations. At 4 wk after treatment, the 1, 10, and
100% concentrations reduced leaf number relative to
the control. Other research showed that allelopathic ac-
tivity from sugarcane leachates did not affect growth of
weed seeds if sown 10 d after leachates were applied,
suggesting microbial degradation, chemical decomposi-
tion, and sorption (Inderjit, 2001b; Sampietro et al.,
2005). Similar to sugarcane, corn (Zea mays L.) residues,
containing vanillic, syringic, p-coumaric, and ferulic
acids, inhibited the shoot growth of corn. On the other
hand, this corn phytotoxicity persisted longer (10 wk)
(Ai-Mezori et al., 1999). It was hypothesized that soil
type would interact with the autotoxicity of the extract,
but there was no significant soil type by concentration
interaction in this experiment (data not shown).
 VIATOR ET AL.: SUGARCANE RESIDUE ALLELOPATHY 1529

Table 2. Comparisons of means for sugarcane growth in a greenhouse as a function of increasing water-soluble extract concentration from
sugarcane postharvest residue for two trials with four replications each averaged across Commerce silt loam and Sharkey clay soils.
Weeks after planting
2 4 6 8 2 4 6 8 8
Concentration Leaf number Plant height Plant dry weight
% no. per plant cm g
0 5.8ab† 7.1a 8.5a 9.4a 27.3a 28.3a 33.7a 46.2a 17.3a
0.1 5.8ab 6.7abc 8.1a 9.6a 27.7a 28.8a 30.6a 46.0a 17.1a
Reproduced from Agronomy Journal. Published by American Society of Agronomy. All copyrights reserved.

1.0 5.3c 6.3c 7.9a 8.9a 26.6a 27.0a 31.4a 44.5a 17.5a
10 5.4bc 6.4c 8.2a 9.5a 26.4a 28.6a 32.7a 45.7a 16.5a
25 5.9a 7.0ab 8.4a 9.6a 28.4a 28.8a 32.0a 47.5a 18.2a
100 5.3c 6.6bc 8.4a 9.4a 23.8a 28.1a 34.7a 46.7a 15.9a
† Means within a column followed by the same lowercase letter are not statistically different using the F probability values and the PROC MIXED macro as
described by Saxton (1998) at a 5 0.05.

On the other hand, there was a significant soil type 3
concentration interaction for the morningglory allelo-
pathy experiment, and thus data were analyzed by soil
type. Morningglory germination was affected only when
seeds were incubated in the Commerce silt loam (Ta-
ble 3). The 0.1, 1, 10, 25, and 100% extract concentra-
tions decreased percentage germination by 24, 40, 38,
53, and 51%, respectively, compared with the control.
At the 25 and 100% concentrations, germination per-
centages were 18% lower than the average of all other
concentrations. Radical growth was affected by extracts
for both soils (Table 3). On the Commerce soil, the 0.1,
1, 10, 25, and 100% extract concentrations decreased
radical length by 6, 7, 7, 10, and 10 mm, respectively,
compared with the control. The 25 and 100% concen-
trations reduced radical length by 3 mm compared with
the mean of all other concentrations. On the Sharkey
clay, the 100% extract concentration reduced radical
length by 5, 4, 5, and 5 mm compared with the control,
0.1, 1, and 10% concentrations. On the Commerce silt
loam, seedling dry weight was reduced by 8 (42%), 11
(58%), and 12 mg (63%) for the 10, 25, and 100%
concentrations relative to the control. The two highest
concentrations decreased weight by 9 mg compared with
the 0.1 and 1% concentrations. Only the 100% con-
centration affected seedling weight on the Sharkey clay.
This treatment reduced weight by 4 (24%) and 5 (28%)
mg relative to the control and the mean of all other
extract concentrations. Previous research indicated that
leachates from sugarcane straw did not affect Ipomoea
species (Lorenzi et al., 1989; Manechini et al., 2005).
Overall, allelopathic activity appeared more pronounced
on the lighter-textured soil compared with the heavy-
textured soil. One possible explanation is that the pH of
the Commerce silt loam was 7.1 compared with 8.0 for
the Sharkey clay. Research has indicated that pH can
affect levels of allelopathic activity (Fujita and Kubo,
2003; Furubayashi et al., 2005).
Postharvest residue extract significantly affected ger-
mination of oat and rye (Table 4). Oat germination was
only affected by the two highest concentrations when
compared with the control, with a 17% reduction re-
corded for the 25 and 100% concentrations. Rye seed
germination was reduced only by the 10, 25, and 100%
concentrations by 12, 12, and 17%, respectively. Radical
growth of oat was reduced by all extract concentrations
by 33 to 60% as compared with the control radicals. The
0.1% concentration increased rye radical growth by 4, 4,
and 5 mm compared with the 10, 25, and 100% con-
centrations. Extracts, though, did not reduce rye radical
growth compared with the control. This contradicts pre-
vious research that demonstrated that benzoic acid from
wheat seedling root exudates inhibited the root growth
of annual ryegrass (Wu et al., 2002). In contrast to oat
and rye, tomato seed germination and radical growth
was not affected by extract treatment. Tomato growth
may not have been affected because it is a dicot unlike
rye and oat. Differential species specificity has been re-
ported with other allelochemicals (Batish et al., 2004).
Previous sugarcane allelopathy research has indicated
inhibitory effects of leachates on root elongation of beg-
gerticks and wild mustard, root growth of lentil, and
seed germination of lettuce (de Carvalho et al., 1996;
Sampietro and Vattuone, 2006a; Singh et al., 2003).

Table 3. Comparisons of means for tall morningglory germination, radical length, and seedling fresh weight when grown in an incubator in
Commerce silt loam and Sharkey clay soil as a function of increasing water-soluble extract concentration from sugarcane postharvest
residue based on two trials with four replications each.
Percentage germination† Radical growth Seedling dry weight
Concentration Commerce Sharkey Commerce Sharkey Commerce Sharkey
% mm mg
0 63a‡ 58a 11a 15a 19a 17b
0.1 39b 54a 5b 14a 19a 17b
1.0 23c 52a 4b 15a 15ab 18ab
10 25c 57a 4b 15a 11bc 20a
25 10d 53a 1c 13ab 8c 17b
100 12d 60a 1c 10b 7c 13c
† Germination percentages, radical lengths, and seedling fresh weight were determined 4 d after dark incubation.
‡ Means within a column followed by the same lowercase letter are not statistically different using the F probability values and the PROC MIXED macro as
described by Saxton (1998) at a 5 0.05.
 1530 AGRONOMY JOURNAL, VOL. 98, NOVEMBER–DECEMBER 2006

Table 4. Comparisons of means for oat, rye, tomato, and sugarcane germination and radical growth of oat, rye, and tomato when grown in
an incubator as a function of increasing water-soluble extract concentration from sugarcane postharvest residue based on two trials with
three replications each.
Percentage germination Radical growth†
Concentration Oat Rye Tomato Sugarcane Oat Rye Tomato
% mm
0 74a‡ 81a 84a 50bc 15a 29ab 12a
0.1 67ab 85a 86a 45bc 9b 31a 12a
1.0 68ab 78ab 88a 65ab 10b 28ab 12a
Reproduced from Agronomy Journal. Published by American Society of Agronomy. All copyrights reserved.

10 62ab 69bc 87a 95a 8b 27b 12a

25 57b 69bc 88a 30c 6b 27b 9a
100 57b 64c 90a 25c 6b 26b 10a
† Radical growth was not measured for sugarcane. Germination percentages and radical lengths were determined 7 d after dark incubation for oat, rye, and
tomato. Germination for sugarcane was determined 14 d after dark incubation.
‡ Means within a column followed by the same lowercase letter are not statistically different using the F probability values and the PROC MIXED macro as
described by Saxton (1998) at a 5 0.05.

Other research has shown that sugarcane leachates
stimulated root growth of a subset of test plants including
pigweed (Amaranthus spp.), radish, sorghum [Sorghum
bicolor (L.) Moench], wheat, and wild mustard at 6 g dry
straw L21 but inhibited growth at higher concentrations.
Synthetic phenolics at 19 mg L21 stimulated the growth of
lettuce, pigweed, arrowleaf sida (Sida rhombifolia L.),
beggerticks, wild mustard, radish, sorghum, and wheat
(Sampietro et al., 2005).
Sugarcane bud germination was increased by 45% by
the 10% extract concentration compared with the con-
trol (Table 4). Statistical analysis for hormetic effects did
reveal that the response curve for sugarcane germina-
tion was due to hormesis (r 2 5 0.82) (Fig. 1). Oat, rye,
and tomato did not exhibit hormesis based on non-
convergence to the log-logistic model (data not shown).
Pearson correlation coefficient responses of sugarcane,
rye, oat, and tomato to the extract were low. Oat, rye,
and tomato had correlation coefficients of 0.26, 0.12, and
350
300
250
percent of control
0.19, respectively; thus, these would not appear to be
good indicator species for sugarcane. A good indicator
species shows correlation coefficients greater than 0.90
with the species under investigation (Ebana et al., 2001).
One of the main reasons these species had poor corre-
lation with the sugarcane response was because of the
hormetic effects demonstrated only with sugarcane.
CONCLUSIONS
Sugarcane postharvest residue showed allelopathic,
autotoxic, and hormetic properties under controlled in-
cubator and greenhouse environments. Benzoic acid was
present in the extracts. Allelopathic activity appeared
more pronounced on the lighter-textured soil compared
with the heavy-textured soil. Due to the hormetic effects
only on sugarcane germination, oat, rye, and tomato were
not good indicator species for sugarcane. Further studies
are needed to establish the impact of benzoic acid in
natural settings.
ACKNOWLEDGMENTS
The authors thank the American Sugarcane League and the
Terrebonne-Barataria National Estuary Program for financial
support to conduct this research.

200
150
100
50
0 Batish, D.R., N. Setia, H.P. Singh, and R.K. Kohli. 2004. Phytotoxicity
0 20 40 60 80 100
Percent concentration of original extract
Fig. 1. Hormetic response of sugarcane bud germination to various
concentrations of postharvest sugarcane residue extracts using the
equation described by Schabenberger et al. (1999): E[Y|x] 5 g 1
{a 2 g/1 1 v exp[bln(x/RD50)]}. E[Y|x] represents the average
response at x dosage, a and g are the upper and lower asymptotes
of the response, v is the initial slope, b is the point of inflection
of the curve, and RD50 is the effective dosage at which 50% of
the total effect is demonstrated. F 5 185.9, p , 0.0001, r 2 5 0.84,
a 5 225, g 5 25, v 5 190, b 5 16, and RD50 5 25.
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