Weed Science Society of America

Quantitative Analysis of Emergence of Seedlings from Buried Weed Seeds with Increasing Soil
Depth
Author(s): Stefano Benvenuti, Mario MacChia and Sergio Miele
Reviewed work(s):
Source: Weed Science, Vol. 49, No. 4 (Jul. - Aug., 2001), pp. 528-535
Published by: Weed Science Society of America and Allen Press
Stable URL: http://www.jstor.org/stable/4046486 .
Accessed: 18/09/2012 09:33

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp

.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact support@jstor.org.

Weed Science Society of America and Allen Press are collaborating with JSTOR to digitize, preserve and
extend access to Weed Science.

http://www.jstor.org
WeedScience,49:528-535. 2001
Quantitative analysis of
buried weed seeds with
Stefano Benvenuti
Corresponding author. Dipartimento di Agronomia
e Gestione dell'Agroecosistema,Universita di Pisa,
Via S.Michele degli Scalzi 2, 56124 Pisa, Italy;
sbenve@agr.unipi.it
Mario Macchia
Sergio Miele
Dipartimento di Agronomia e Gestione
dell'Agroecosistema,Universita di Pisa, Via
S.Michele degli Scalzi 2, 56124 Pisa, Italy
Knowledge of weed biology has aroused increasing inter-
est in recent years in the context of weed management strat-
egies (Bhowmik 1997). Such knowledge is essential to max-
imize the effectiveness of agronomic practices (Forcellaet al.
1993) and to simulate weed dynamics. The greater insight
into weed dynamics gained through this approach has
opened up new horizons in crop protection, allowing ratio-
nal utilization of chemical control. The overall aim is to
apply herbicides exclusively, and proportionately, only in
cases of genuine risk of weed-crop competition.
Weed dynamics simulations have considered a number of
patterns of the weed life cycle, including agronomic soil
disturbance (Ball 1992; Mohler 1993), response to ecolog-
ical and climatic factors (Alm et al. 1993; Forcella 1993,
1998), and even subsequent weed growth within crops
(Kropf and Van Laar 1993). Crucial to successful simulation
is knowledge of the extent of the soil seed bank (Forcella
1992; Roberts and Ricketts 1979; Zhang et al. 1998), an-
nual seed production (Benvenuti et al. 1994; Cardina and
Norquay 1997), and aspects of buried seed ecology (Pons
1991) that can influence seed longevity (Lueshen and An-
dersen 1980). Ecological factors have been shown to play a
major role, not only by influencing secondary dormancy
(Baskin and Baskin 1985) but also by either inducing or
inhibiting germination. Thus, it is well known that light
(Ballare et al. 1992), temperature (Benvenuti and Macchia
1993), soil water content (Roberts and Potter 1985), and
degree of soil compaction (Parejaand Staniforth 1985) rep-
resent the main factors limiting buried seed germination.
528 * Weed Science 49, July-August 2001
emergence of seedlings from
increasing soil depth
Trialswere carriedout to investigatethe effectsof seed burialdepth on seedling
emergencerateof 20 weed species.Markeddepth-mediatedvariationin emergence
abilityof the differentspecieswas observed,togetherwith a generalpatternof de-
creasingemergencewith increasingsoil depth.At 10 cm, only johnsongrass, velvet-
leaf,catchweedbedstraw,and cutleafgeraniumemerged,albeitonly in limitednum-
bers. Speciesmost severelyinhibitedby burialdepthwerebuckhornplantain,large
crabgrass,common purslane,chickweed,and corn spurry,none of which emerged
from beyond 6 cm. In all species,depth-mediatedinhibitionwas found to be sig-
moidal (polynomialregression).In addition,the numberof seedlingsand rate of
seedlingemergencedecreasedwhen depth of burialincreased.The depth at which
the numberof emergedseedlingswas halvedvariedby speciesand rangedfrom3.6
cm for common purslaneand chickweedto 7 cm for velvetleafand catchweed
bedstraw.Excessiveburialdepth generallyinduceddormancy(in roughly85% of
cases)ratherthan suicidegermination.A close inverserelation(second-degreeequa-
tion) betweenseed unit weight and depth-mediatedinhibitionwas observed.The
physiologicalinvolvementof depth inhibitionin seed bankecologyis discussed.
Nomenclature: Buckhornplantain,PlantagolanceolataL. PLALA;catchweedbed-
straw, GaliumaparineL. GALAP;common chickweed,Stellariamedia(L.) Vill.
STEME;common purslane,PortulacaoleraceaL. POROL;corn spurry,Spergula
arvensL. SPRAR;cutleafgeranium,Geraniumdissectum L. GERDI;johnsongrass,
Sorgumhalepense(L.) Pers.SORHA;largecrabgrass,Digitariasanguinalis(L.) Scop.
DIGSA;velvetleaf,Abutilontheophrasti
MedicusABUTH.
Key words: Seed germination,seedlingemergence,burialdepth,soil seed bank.
Germination is also known to be inhibited by soil depth
(Holm 1972; Stoller and Wax 1973). The biological reason
for depth inhibition has not yet been fully clarified. Previous
studies suggest this may not be due merely to lack of light
(Benvenuti 1995), but also to decreasingthermal fluctuation
with increasing burial depth, as thermal fluctuation consti-
tutes a known germination trigger (Roberts and Totterdell
1981). An alternative or possibly complementary explana-
tion is based on gas diffusion, which is inversely correlated
with burial depth. Reduced gas exchange is believed to be
capable of inducing secondary dormancy (Benvenuti and
Macchia 1995). Presence of CO2 deriving from soil biolog-
ical activity is probably also involved in this complex mech-
anism (Karssen 1982). It is therefore of prime importance
to acquire knowledge on weed seed distribution in soil
(Cousens and Moss 1990; Grundy et al. 1996) and on the
emergence ability of various weed species (Alm et al. 1993).
Emergence from different soil depths has been found to be
proportional to seed energy reserves (Lafond and Baker
1986). It is also influenced by the typology of such reserves,
depending on the amount of oxygen required for their uti-
lization (Raymond et al. 1985). Energy reservesare vital for
seedling growth prior to emergence, because given the ab-
sence of light, growth takes place completely autotrophically.
Furthermore, independent of the physiological aspects char-
acterizing this agronomically important stage, weed species
may differ in their ability to emerge from the various soil
layers.
Accurate knowledge of these parameters exerts a marked
influenceon the degreeof precisionof weed dynamicsim-
ulations.It consequentlyaffectssuccessfulagronomicappli-
cation of weed dynamicsmodelingand, thus, the possibility
of setting up rationalweed controlstrategies.
The purposeof this workwas (1) to performquantitative
analysisof the seedlingemergencecharacteristicsof 20 dif-
ferentweed speciesas a function of depth of burialand (2)
to investigatepossiblecorrelationsbetweenemergenceabil-
ity and seed size.
Materials and Methods
Trialswere carriedout at the Seed Researchand Testing
Station(InternationalSeedTestingAssociationapproved)of
the AgronomyDepartmentof Pisa University,Italy.Seeds
of 20 weed species-prostrate knotweed (Polygonumla-
phathifoliumL. POLLA),redrootpigweed (Amaranthus re-
troflexusL. AMARE),buckhornplantain,jimsonweed(Da-
tura stramoniumL. DATST), Canadathistle [Cirsiumar-
vense(L.) Scop. CIRAR], large crabgrass,blackgrass(Alo-
pecurus myosuroides Huds. ALOMY), johnsongrass,
barnyardgrass [Echinochloacrus-galli(L.) Beauv.ECHCG],
wild mustard[Brassicakaber(DC.) L.C. WheelerSINAR],
common lambsquarters(Chenopodium albumL. CHEAL),
black nightshade(SolanumnigrumL. SOLNI), velvetleaf
(Abutilontheophrasti),hairybittercress(CardaminehirsutaL.
CARHI), catchweedbedstraw,cutleaf geranium,common
purslane,field bindweed(Convolvolus arvensisL. CONAR),
common chickweed,and corn spurry-were collectedat full
ripeningin spring(une 15) and summer(August25) 1998
near Pisa, Italy.Seeds were cleanedimmediatelyafter har-
vesting and placed in hermeticallysealedscrew-topplastic
jars.Seedswerestoredin darknessat room temperature(20
C) and relativehumidity < 10%, determinedaccordingto
ISTA methods (1999) based on the loss in weight when
seedsaredrieduntil use in September1999. Beforeseeding,
18 weed specieswerestratifiedat 4 C for 2 wk to overcome
any dormancy.An exceptionto this procedurewas adopted
for velvetleafand field bindweed(Horowitzand Taylorson
1985; Mitich 1991, respectively),whichwereburied(in jute
bags)in the field 30 cm deep for 1 yr priorto use (Septem-
ber 1998-September1999) to overcomecharacteristic seed
hardness,according to a previouslydescribedprocedure
(Cardinaand Sparrow1997).
Seeding
Fifty seeds of each of the 20 weed specieswere sown in
plasticpots (15 by 15 by 25 cm) filledwith a silt-loamsoil
(typic xerofluvent soil; 23% sand, 52% silt, 25% clay). Soil
was obtainedby excavationsfrom a depth of over 1 m in
orderto avoid the presenceof preexistingseeds capableof
falsifyingthe real experimentaldata of emergencerates.In
spite of obtainingsoil froma deep soil layer,the relativesoil
characteristicswere similar to a normal topsoil (pH 6.8,
0.08% total N, 21 mg kg-1 availableP205, 265 mg kg-l
exchangeableK20) and no particularproblembasedon aer-
obic or anaerobicconditions was found. Pots were filled
gravimetricallywith the soil and packed with a uniform
strength to avoid differential resistance to seedling emer-
gence. Seeding depths were 0, 2, 4, 6, 8, 10, or 12 cm. The
study was a completely randomized design with three rep-
licates for each seeding depth and the study was repeated.
Seedson the soil surfacewere coveredwith moistenedfilter
paper1at first (12 h) to stimulateimbibition.
Buried Seed Incubation
Seeded pots were placed in climate-controlledcabinets
presetto alternatingtemperaturesof 25/30 C for predom-
inantly summer weeds (prostrateknotweed, redroot pig-
weed, jimsonweed,Canadathistle,largecrabgrass,johnson-
grass, barnyardgrass, common lambsquarters,black night-
shade, velvetleaf,field bindweed, and common purslane)
and 15/20 C for predominantlyspring weeds (buckhorn
plantain,blackgrass,wild mustard,hairybittercress,catch-
weed bedstraw,cutleafgeranium,common chickweed,and
corn spurry).The same 12-h dark/lightphotoperiodwas
maintainedfor both temperatures.Light intensityof 100
Lmol m-2 s-1 was produced by cool fluorescent tubes2 and
was measuredwith a spectroradiometer.3 During incuba-
tion, pots were moistenedby subirrigation.
Seedling Emergence
Emergedseedlingswere counted daily at cotyledonap-
pearanceand removed.Seedlingcounts were stopped3 to
5 d afterno moreemergencewas recorded.Meanemergence
time (MET) was calculatedas
MET = l(n x g)lN,
where n is the numberof seedlingsemergingper day,g is
the number of days needed for emergence,and N is the
total numberof emergedseeds.Foreachseedingdepth,per-
centageof soil depth inhibitionwas calculatedfor eachspe-
cies as a function of unburiedseed germination(0% inhi-
bition).
Ungerminated Seed Recovery and Germination
Test
After emergencetests, soil was removedfrom pots and
washed to determinethe fate of ungerminatedseeds (dor-
mant and/or germinatedwithout emergence).A fine metal
sieve (400 gm) was used for seed recovery.Suicidegermi-
nation (not followedby seedlingemergence)was calculated
as the differencebetweentotal seedsand the sum of recov-
ered seedsand emergedseedlings.Fifty ungerminatedseeds
by each pot from the deep burialof 12 cm were imbibed
with 3 ml of distilledwateron a single sheet of filterpaper
placedin 6-cm petri dishes.Germinationtestswere carried
out in the same photothermicconditionsas the emergence
tests.
Seed Weight
Seed weight was determinedby weighing 1,000 seeds
chosen randomlyaccordingto ISTA rules for seed testing
(ISTA 1999).
Calculation of Depth of 50% Emergence
Inhibition
Polynomial regressions were calculated that showed the
best fit of the biological response of weed seed emergence-
inhibition at increasing soil depths. These polynomial equa-
Benvenutiet al.: Buriedweed seedlingemergence * 529
 100 - y--0.121 xi .92xA2.61 x-0.3 - y-0.038x4-1 31i &1875x-2.9 100
R20.98 > R2=0.98 9
80 so~~~~~~~~~~~~~~~~~~~~~~~~8

Prostrate knotweed Redroot pigweed

40 40

y--0.024x-o027 1
+5.28x-3 4 -
_,0 j2 09x+03 100

so
3.85
b so8
60 1 i/ t z
Jimsonweed
~~~~Buckhorn
plantain
100 - _=016Y25I00 2=ooe -Fi~ x7.9x-03 10

6.1~~~~~~~~~~~~~~,

a +
r20Xta
e-N20da /
00 eoL g lCda ethistle Large crabgrass e a C

LI~~~~~~R-09 R -0.98

:r,ef 0
'|~~~~~~~~~~~~~~~~~~~~~~-.
,
[0 1 k 05260 6x+5.20-6 - y--0.l1 4x-2,45x-4,84+1 13 ? i100 ? uz

3o / 80
C:so- sor

o-a b| /2 4 1
Wi usad B
io L L a_a d as L2

o b Caad thitl 10 12
0)~~~ Larg crabgrass

100 - y--=Ol 73x+2,57x3-1 S x - y=-0097x -1 1.34x1 BE io0

Seeding depth (cm) Seeding depth (cm)

FIGURE 1. Seedling emergence and relative degree of depth inhibition (as percentage of soil surface germination) of the 20 selected weed species as a
function of increasing burial depth. Means followed by the same letter do not differ at P < 0.05 according to the Student-Newman-Keuls test. The
equations of inhibitions (significant for P < 0.01) and the corresponding R2 values are reported. Arrows indicate the values of 50% depth inhibition.

tions of soil depth inhibition activity were used to identify
soil depths capable of reducing emergence to 50% of soil
surface germination. Depth values were plotted with the
corresponding 1,000-seed weight and fitted with a quadratic
polynomial regression.
Statistical Analysis
The two experiments did not differ in seed response and
data were pooled. After testing for homogeneity of variance,
arcsine transformation of emergence percentages was nec-
essary.Angular values were subjected to analysis of variance
(ANOVA) using the Student-Newman-Keuls test (P <
0.05) for means separation. Inhibition data of each species
tested were fitted by the corresponding polynomial regres-
sion that adequately described the biological response of
weed seed germination and emergence. These equations
gave the soil depths at which emergence rates reached 50%
by using a modified "x-intercept"method (Wiese and Bin-
ning 1987). The intercept between the polynomial regres-
sion and the translated x-axis on the selected y for 50%
depth inhibition shows the relative soil depth inhibition for

530 * Weed Science 49, July-August 2001

 100 -Y-0.152A2.30x2.58x-3.4 , y--0.170X+3.56x-3.C66 100

ff
80~~~ Common lambsquarters| Black nightshade

R2-0.97 Rf -0.98 >

so0 a so
e01 a x,, < d d | 1 /8 X e e 2

100 _y=-0.149xt-2,gBx-5,87x-i-08
- =OeOt 1fh i .8lO21x-3
0 c _ 100
40 -
e
~~~~~~~~~~~~~~~~~~~~~~40

b Velvtle=f

E
20 a ~~~~~~~~~~~~~~~~~~~~~~~~d20
b / L b 3QS / 80

50
2 [
tom~~~mmon Blcknihshde;
rlambqurtr

so~~~~~~~~~~~~~~~~~~~5
a

SX a
150 -a
20L
50 ,~~~ Common7.
e
chcked
t-d-d e d
e g or3pury
e e
5
2

40 ~~~~~~~~~~~~~~~~~~~~~~~~40

50 60
20~~~~~~~~~~~~~~~~~~~~~~~~~~
2 I4 87 10 18
68 i Con mon Cachicweed 1 01
OKO eedigdpt cm eeigdet Cutleafor
gpuranium
(cm)C
FIGURE 1. Continued. 7-1 1~, 1

FIGURE_1. Continued.

Commonept
purian bedn Fieldhbinwee

each species. For each statistical analysis, commercial soft-
ware (CoStat4) was used.
Results and Discussion
Figure 1 shows the emergence of the 20 weed species
studied as a function of depth of burial. Emergence de-
creased with increasing burial depth, with 12 cm represent-
ing the limit of total inhibition at which no species was able
to emerge. For each of the species examined, depth inhibi-
tion was found to be highly significant (P < 0.01) by fitting
the relevant data by polynomial regression.
Analysis of the outcome for individual species (Table 1)
shows that prostrate knotweed, redroot pigweed, and jim-
sonweed had emergence less than 10% at a burial depth of
8 cm, whereas buckhorn plantain showed complete inhibi-
tion at this depth. Large crabgrasswas unable to emerge
from a depth greater than 6 cm, whereas johnsongrass did,
albeit at a minimum percentage (roughly 5%), from as deep
as 10 cm. However, for the majority of species such as Can-
ada thistle, blackgrass,barnyardgrass,common lambsquart-
ers, black nightshade, and wild mustard, the depth limit was
considerably less then 10 cm, and emergence was very low
(ranging between 5 and 10%) even at a burial depth of 8
cm. The greatest ability to maintain emergence with increas-
ing depth, although at low levels, was shown by velvetleaf,

Benvenuti et al.: Buried weed seedling emergence * 531

TABLE 1. Effect of seeding depth on mean emergence time of the 20 tested weed species. (Means are followed by the relative standard
errors.)
Meanemergencetime (seedingdepth,cm)b
Weedspeciesa 0 2 4 6 8 10 12
d
Prostrate knotweed 6.6 (1.3) 8.2 (1.6) 10.5 (1.9) 12.2 (2.5) 16.2 (2.9)
Redroot pigweed 4.3 (1.1) 5.7 (1.1) 7.5 (1.2) 9.5 (0.8) 14.7 (2.3)
Buckhorn plantain 7.4 (1.3) 8.9 (0.5) 11.6 (0.9) 13.2 (1.1)
Jimsonweed 7.2 (1.1) 8.8 (0.4) 11.4 (0.8) 12.9 (1.0) 21.9 (3.7)
Canada thistle 5.1 (0.7) 6.5 (0.3) 8.5 (0.7) 10.7 (0.9) 16.5 (2.2)
Large crabgrass 6.0 (0.6) 7.4 (0.3) 9.6 (0.8) 12.4 (1.0)
Blackgrass 5.5 (0.4) 6.8 (0.2) 8.8 (0.2) 11.3 (0.9) 17.1 (2.1)
Johnsongrass 6.3 (0.3) 7.3 (0.3) 9.4 (0.4) 12.1 (1.3) 16.7 (2.4) 20.8 (3.1)
Common lambsquarters 6.7 (0.3) 7.9 (0.4) 10.2 (0.5) 13.3 (1.5) 19.5 (3.0)
Black nightshade 7.3 (0.5) 8.5 (0.6) 12.0 (0.6) 15.8 (1.9) 23.2 (3.7)
Barnyardgrass 6.1 (0.4) 7.2 (0.5) 9.3 (0.4) 12.4 (1.7) 17.6 (3.1)
Wild mustard 3.7 (0.3) 4.8 (0.4) 6.2 (0.3) 8.4 (0.8) 11.1 (2.2)
Velvetleaf 5.6 (0.6) 6.8 (0.7) 8.8 (0.9) 11.3 (1.0) 16.5 (3.2) 19.7 (2.9)
Hairy bittercress 4.1 (0.5) 5.8 (0.6) 7.5 (0.8) 10.5 (0.9) 12.7 (2.7)
Catchweed bedstraw 6.8 (0.7) 7.7 (0.8) 10.1 (1.0) 13.2 (1.2) 19.1 (2.9) 22.6 (3.2)
Cutleaf geranium 7.3 (0.8) 8.5 (0.9) 11.0 (1.2) 13.6 (1.3) 20.2 (3.0) 23.1 (3.4)
Common purslane 5.9 (0.5) 7.2 (0.8) 9.4 (1.1) 12.2 (1.4)
Field bindweed 6.4 (0.7) 7.8 (1.0) 10.1 (1.3) 13.2 (1.6) 19.2 (2.7)
Common chickweed 5.7 (0.6) 7.1 (1.0) 9.2 (1.4) 12.0 (1.8)
Corn spurry 6.3 (0.5) 7.8 (1.2) 10.1 (1.6) 13.2 (2.1) - - -
a Barnyardgrass,Echinochloa crus-galli; black nightshade, Solanum nigrum; blackgrass, Alopecurusmyosuroides,buckhorn plantain, Plantago lanceolata;
Canadathistle, Cirsium arvense,catchweed bedstraw, Galium aparine; commonchickweed, Stellaria media; commonlambsquarters, Chenopodiumalbum;
common purslane, Portulaca oleracea;corn spurry, Spergula arvensis, cutleaf geranium, Geranium dissectum;field bindweed, Convolvolusarvensis, hairy
bittercress, Cardaminehirsuta;jimsonweed, Datura stramonium;johnsongrass, Sorghumhalepense;large crabgrass,Digitaria sanguinalis,prostrateknotweed,
Polygonumlaphathifolium;redrootpigweed, Amaranthusretroflexus-,velvetleaf, Abutilon theophrasti;wild mustard, Brassicakaber.
b Missing values indicate no seedling emergence.

catchweed bedstraw, cutleaf geranium, and field bindweed,
which were generally able to germinate and reach the soil
surface even from 10 cm (5 to 15%). In contrast, hairy
bittercress, common purslane, common chickweed, and
corn spurry showed marked inability to emerge from deep
burial and a limit of about 6 cm.
In all species tested, emergence decreased slightly with
shallow burial (2 cm) and then decreased exponentially at
greater depths. However, some authors (Chancellor 1964;
Mohler and Galford 1997; Wiese and Davis 1967), who
performed similar experiments on analogous weed species,
reported a favorable effect of slight burial (roughly 0.5 to 1
cm) on emergence. Such an effect was not detected in the
present study. This could have been because of water stress
near the soil surface in the cited studies, a phenomenon that
was avoided in the present test by covering the soil surface
with filter paper. Additionally, in our study, the first depth
below the surface layer studied was 2 cm, a depth that did
not include the zone in which variable responses were de-
scribed in the cited studies.
Other factors leading to difficulty in comparing results
obtained in this study with those reported by other authors
include possible differences in dormancy characteristicsof
the various species used, the particular temperature and hu-
midity conditions, and above all, the different soil charac-
teristics. Soil particle size has been shown to influence soil
physical characteristics,which can interferewith buried seed
germination and emergence (Cussans et al. 1996). Despite
these general differences, there is overall agreement that
emergence inhibition increases proportionately with depth
of seed burial in soil (Benvenuti and Macchia 1997; Black-
shaw 1992; Mester and Buhler 1991; Vleeshouwers 1997;
Webb et al. 1987).
In our study, burial depth also exerted marked influence
on MET in each of the species tested (Table 1). Each 2 cm
increase in distance below the soil surface resulted in a sig-
nificant (P < 0.05) increase in emergence time. This delay
in emergence from deeper soil levels means that deeply bur-
ied seeds have a disadvantage in competition with crops.
Use of equations to representinhibition induced by burial
depth (expressedas a percentage compared to unburied seed
germination) allowed identification of the burial depth at
which emergence was halved. This depth varied depending
on the species examined, ranging from 3.6 cm in common
purslane and common chickweed to 7 cm with velvetleaf
and catchweed bedstraw. Along this gradient, Canada this-
tle, black nightshade, and barnyardgrassshowed depth-me-
diated emergence (5.3 cm) approaching mean 50% emer-
gence values of the 20 species tested (mean 5.1 cm). The
level of emergence we found was slightly greaterthan values
detected in field conditions by other authors (reviewed by
Mohler 1993). This discrepancy could be due to the greater
physical constriction in a field with undisturbed soil than in
our dry-sieved experimental soil. It has been shown that soil
compaction impairs the ability of seeds to reach the soil
surface during preemergence seedling growth (Hegarty and
Royale 1978). Furthermore,soil compaction can act directly
on seeds by limiting germination (Pereja and Staniforth
1985) or even by inducing dormancy (Terpstra1995). How-
ever in our study, the effects of soil compaction were not
examined. A relative comparison was made of depth-medi-
ated emergence among different weed species under similar,

532 * Weed Science 49, July-August 2001

 100 100
T T

a, 9070
T
E T T IT T T T T0 90 :
0
_~~~~~
U)
Tz3 + ~~ si g8
- ,
20J |80 T
T 11?T?*
? 8 v

0 PL IU.
Q 70
'C
cn ?70~u)
E -
60 Q )
0
60
~C, SD 0
UD 50 - 0
cn E
-o 0 -
0 U,0Q-
~~~~~ 30 ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ -3Q0
cn 20 - ~~~ U)
~~~~~~20
10
0 0
POLLA PLA CI;AR ALOMY ECHCG CHEAL 'ABUh GAAPPOROL SEM
AWAE DATST DIOSA SORHA SINA SOLNI CARHI GERDI CONA SPRA

Weed species
FIGURE 2. Percentage of suicide germination (as percentage of seeded seeds) and relative percentage of ungerminated seeds retrieved after the emergence
test from the greatest (12 cm) burial depth. The means across species are indicated by the solid line. Vertical bars indicate standard errors of the mean.

noncompacted soil conditions. Examination of seeds recov- ed by inducing depth-mediated dormancy. Dormancy
ered from a burial depth at which none of the seeds of the mechanisms acquired by weeds through evolution would in-
20 species tested succeeded in emerging showed that almost clude perception of excessive seed burial depth (Mapes et al.
all (approximately 85%) remained completely dormant (Fig- 1989). This speculation was confirmed by noting that res-
ure 2), so that germination was very limited (only 15%). toration of optimal incubation conditions in petri dishes
These findings suggest that seeds of the various species (data not shown) proved sufficient for all 20 species tested
perceived unfavorable germination conditions and respond- to achieve germination percentages similar to values ob-
served in unburied seeds. Depth-mediated dormancy was
reported by Wesson and Wareing (1969) as early as 1969,
TABLE 2. Weed species (listed as a function of decreasingsize, and although it has not yet been fully clarified, it appears
1,000-seed weight) tested in the trials. Means are followed by their to be linked to increasing difficulty in gas exchange with
relative standard errors.
increasing soil depth. In particular, induction of seed dor-
Weed species 1,000-seedweight mancy seems to be mediated by lack of 02 (Benvenuti and
Macchia 1995, 1998) or by an increase in CO2 (Holm
g
1972) deriving from seed metabolism.
Catchweed bedstraw 11.41 + 0.143
Field bindweed 9.777 ? 1.034
However, the germination behavior of seeds buried at in-
Velvetleaf 8.645 ? 0.763 creasing depths may also be linked to seed energy reserves,
Jimsonweed 8.192 + 0.847 because reserves are known to be crucial in some crop spe-
Johnsongrass 5.115 ? 0.574 cies to allow activation of seed metabolism even under ox-
Cutleaf geranium 2.133 ? 0.312 ygen deficiency (Al-Ani et al. 1985). It should not be over-
Blackgrass 1.858 + 0.911 looked that virtually the entire seed bank is subjected to
Wild mustard 1.813 + 0.203 some degree of hypoxia, given that oxygen concentration is
Canada thistle 1.697 + 0.175 inversely correlated with depth (Drew 1990). Seeds germi-
Prostrateknotweed 1.603 ? 0.234
Barnyardgrass 0.885 + 0.075
nating at excessive depth would have little chance of reach-
Black nightshade 0.803 ? 0.094 ing the soil surface that represents the target of the energy-
Buckhorn plantain 0.750 + 0.083 limited autotrophic phase (beginning of photosynthesis).
Large crabgrass 0.508 ? 0.052 In this perspective, seed specific weight could play a major
Common lambsquarters 0.490 + 0.052 role in allowing germination at a considerable distance be-
Redroot pigweed 0.424 + 0.051 low the soil surface. Table 2 shows the specific weight (re-
Common chickweed 0.373 ? 0.041 ferring to 1,000 seeds) of each of the 20 species studied.
Corn spurry 0.231 ? 0.034 Noticeable variation was found, ranging from over 11 g in
Common purslane 0.092 + 0.008
Hairy bittercress 0.088 ? 0.007 catchweed bedstraw to a more than 1,000-fold lower seed
weight in hairy bittercress (both less than 0.1 g). Among
a Barnyardgrass,
Echinochloacrus-galli;
blacknightshade,Solanumnigrum; intermediate values, specific weights in the upper range were
blackgrass,Alopecurusmyosuroides,buckhorn plantain,Plantagolanceolata;
Canadathistle, Cirsiumarvense-,catchweedbedstraw,Galiumaparine,com- recorded for field bindweed (9.7 g), velvetleaf (8.6 g), and
mon chickweed,Stellariamedia;commonlambsquarters, Chenopodiumal- jimsonweed (8.2 g) and in the lower range for redroot pig-
bum;commonpurslane, Portulacaoleracea; cornspurry,Spergulaarvensis, weed and common lambsquarters (both slightly below 0.5
cutleafgeranium, Geranium dissectum;fieldbindweed, Convolvolus
arvensis, g).
hairybittercress,Cardamine hirsuta;jimsonweed, Daturastramonium; john-
songrass, largecrabgrass,
Sorghumhalepense-, prostrate
Digitariasanguinalis, A possible correlation between seed specific weight (Table
knotweed,Polygonumlaphathifolium; redrootpigweed,Amaranthusretro- 2) and depth-mediated 50% emergence inhibition was ex-
flexus-,velvetleaf,
Abutilontheophrasti;wildmustard, Brassicakaber. plored (Figure 1). A statistically significant correlation (P <

Benvenuti et al.: Buried weed seedling emergence * 533


C) 4-1 .-
7My
a 30
3456
C 08
0
0)
eq ( y=- 0.036 x2he0.639
o o R 0.75
FIUE 3. Rersinbtenbra0et eesr ordc
- 4-- 0
n 0 1 2 3 4 5 6 7 8
1,000 seed weight (g)
FIGuRE 3. Regression between burial depth necessary to reduce emergence
of the 20 weed species tested to 50% and corresponding 1,000-seed weight.
The equation (significant for P > 0.05) and the corresponding R2 value
are shown.
0.05) according to a second-degree equation was observed
(Figure 3). This indicates that seed weight can represent a
valid tool, at least in the species tested, for identifying the
emergence profile of the active soil seed bank (i.e., the like-
lihood of emergence and consequent crop invasion).
A relation between seed size and emergence has been re-
ported for various crop species (Harper and Obeid 1967;
Heather and Sieczka 1991; Taylor and Ten Broeck 1980;
Weaver 1980) and for wild species (Bond et al. 1999). But
because such trials were conducted in the field, it was not
possible to distinguish between suicide germination and seed
dormancy as causes of failed emergence. The present labo-
ratory tests demonstrated that failed emergence was mainly
due to depth-imposed dormancy. This mechanism of ger-
mination inhibition appears to be an important survival
strategy, allowing seed bank perpetuation. It could underlie
the pronounced seed longevity (Burnside et al. 1996) that
leads to a persistent seed bank (Thompson 1987).
In addition, a relation between seed longevity and unit
weight has been suggested (Thompson et al. 1993). It has
been hypothesized further that the pronounced fluctuation
in seed unit weight sometimes present even within the same
species (Milberg et al. 1996) could favor depth-mediated
dormancy heteroblasty of buried weed seeds. This would
ensure that buried dormant seeds maintain their germina-
tion ability despite agronomic disturbances such as appli-
cation of herbicides, tillage, or harvesting, suggesting that
weed seed size may be correlated with resistance to unfa-
vorable ecological situations (Hodkinson et al. 1998). It is
worth pointing out that a seed unit weight effect has been
described in curly dock (Rumex crispusL.) and broadleaf
dock (Rumex obtusijfoliusL.), where seed size was shown to
influence emerged seedling competitiveness (Cidecydan and
Malloch 1982).
Results obtained in this study are of interest for two rea-
sons. First, they clarify the complex ecophysiology of soil
depth inhibition of weed seed germination. Second, they
demons.traqtethat, withi few excepntions- (jonsonngrass, vel-
534 * Weed Science 49, July-August 2001
that involve or simulate seed bank dynamics. Only by
achievingprecisionin seed dynamicsimulationwill it be-
come possible to make weed control optimally effective
0 while minimizingunjustifiedherbicideapplication.This will
favor harmonizationof ecologicalconcernswith the eco-
nomic advantagesexpectedfrom modernagriculture.
Sources of Materials
x+ 4.12
1 Filter paper, Watchman No. 1. Fisher Scientific, Pittsburgh,PA
15219.
mrec 2 Fluorescent lamp, Philips TLF 20W/33, Eindhoven,
The
Netherlands.
9 10 11 12 3 Spectroradiometermodel 1800. Licor, Inc., Lincoln, NE.
4 CoHort Software, Minneapolis, MN 55419.
Literature Cited
Al-Ani, A., F. Bruzau, P. Raymind, V. Sain-Ges, J. M. Leblank, and A.
Pradett. 1985. Germination, respiration and adenylate charge of seeds
at various oxygen pressures. Plant Physiol. 79:885-890.
Alm, D. M., E. W Stoller, and L. M. Wax. 1993. An index for predicting
seed germination and emergence rates. Weed Technol. 7:560-569.
Ball, D. A. 1992. Weed seedbank response to tillage, herbicides, and crop
rotation sequence. Weed Sci. 40:654-659.
Ballare, C. L., A. L. Scopel, R. A. Sanchez, and S. R. Radosevich. 1992.
Photomorphogenic processes in the agriculturalenvironment. Photo-
chem. Photobiol. 56:777-788.
Baskin, J. M. and C. C. Baskin. 1985. The annual dormancy cycle in
buried weed seeds: a continuum. Bioscience 35:492-498.
Benvenuti, S. 1995. Soil light penetration and dormancy of Jimsonweed
(Datura stramonium)seeds. Weed Sci. 43:389-393.
Benvenuti, S. and M. Macchia. 1993. Calculation of threshold temperature
for the development of various weeds. Agric. Mediterr. 123:252-256.
Benvenuti, S. and M. Macchia. 1995. Hypoxia effect on buried weed seed
germination. Weed Res. 35:343-351.
Benvenuti, S. and M. Macchia. 1997. Germination ecophysiology of bur
beggarticks (Bidens tripartita) as affected by light and oxygen. Weed
Sci. 45:696-700.
Benvenuti, S. and M. Macchia. 1998. Phytochrome mediated germination
control of Datura stramoniumL. seeds. Weed Res. 38:199-205.
Benvenuti, S., M. Macchia, and A. Stefani. 1994. Effects of shade on re-
production and some morphological characteristicsof Abutilon theo-
phrasti Medicus, Datura stramoniumL. and SorghumhalepenseL. Pers.
Weed Res. 34:283-288.
Bhowmik, P. C. 1997. Weed biology: importance to weed management.
Weed Sci. 45:349-356.
Blackshaw, R. E. 1992. Soil temperature, soil moisture and seed burial
depth effects on redstem filaree (Erodiumcicutarum)emergence. Weed
Sci. 40:204-207.
Bond, W. J., M. Honig, and K. E. Maze 1999. Seed size and seedling
emergence: an allometric relationship and some ecological implica-
tions. Oecologia 120:132-136.
Burnside, 0. C., R. G. Wilson, S. Weisberg, and K. G. Hubbard. 1996.
Seed longevity of 41 weed species buried 17 years in eastern and
Western Nebraska. Weed Sci. 44:74-86.
Cardina, J. and H. M. Norquay. 1997. Seed bank production and seedbank
dynamics in subthreshold velvetleaf (Abutilon theophrasti)populations.
Weed Sci. 45:85-90.
Cardina, J. and D. H. Sparrow. 1997. Temporal changes in velveatleaf
(Abutilon theophrasti)seed dormancy. Weed Sci. 45:61-66.
Chancellor, R. J. 1964. Emergence of weed seedlings in the field and the
effects of different frequencies of cultivation. Pages 599-606 in Pro-
ceedings of the Seventh British Weed Control Conference, Brighton.
Cidecydan, M. A. and A.J.C. Malloch. 1982. Effects of seed size on the
germination, growth and competitive ability of Rumexcrispusand Ru-
mex obtusifolius.J. Ecology 70:227-232.
Cousens, R. and S. R. Moss. 1990. A model of the effects of cultivation
on the vertical distribution of weed seeds within the soil. Weed Res.
30:61-70.
Cussans, G. W., S. Raudonius, P. Brain, and S. Cumbenworth. 1996. Ef-
fects of depth of seed burial and soil aggregate of Alopecurusmyosu-
 roides, Galium aparine, Stellaria media and wheat. Weed Res. 36:133-
141.
Drew, M. C. 1990. Sensing soil oxygen. Plant Cell Environ. 13:681-693.
Forcella, F. 1992. Prediction of weed seedling densities from buried seed
reserves.Weed Res. 32:29-38.
Forcella, F. 1993. Seedling emergence model for velvetleaf. Agron. J. 85:
929-933.
Forcella, F. 1998. Real-time assessment of seed dormancy and seedling
growth for weed management. Seed Sci. Res. 8:201-209.
Forcella, F., K. E. Oskoui, and S. W. Wagner. 1993. Application of weed
seed bank ecology to low-input crop management. Ecol. Appl. 3:74-
83.
Grundy, A. C., A. Mead, and W Bond. 1996. Modeling the effect of weed-
seed distribution in the soil profile on seedling emergence. Weed Res.
36:375-384.
Harper, J. L. and M. Obeid. 1967. Influence of seed size and depth of
sowing on the establishment and growth of varieties of fiber and oil
seed flax. Crop Sci. 7:527-532.
Heather, D. W. and J. B. Sieczka. 1991. Effect of seed size and cultivar on
emergence and stand establishment of broccoli in crusted soil. J. Am.
Soc. Hortic. Sci. 116:946-949.
Hegarty, T. W. and S. M. Royale. 1978. Soil impedence as a factor reducing
crop seedling emergence, and its relation to soil conditions at sowing,
and to applied water. J. Appl. Ecol. 15:897-904.
Hodkinson, D. J., A. P. Askew, K. Thompson, J. G. Hodgson, J. P. Bakker
and R. M. Bekker. 1998. Ecological correlates of size in the British
flora. Funct. Ecol. 12:762-766.
Holm, R. E. 1972. Volatile metabolites controlling weed germination in
soil. Plant Physiol. 50:293-297.
Horowitz, M. and R. B. Taylorson. 1985. Behaviour of hard and permeable
seeds of Abutilon theophrastiMedic. (Velvetleaf). Weed Res. 25:363-
372.
[ISTA]. 1999. International rules for seed testing. Seed Sci. Technol.
27(Suppl.):50-52.
Karssen, C. M. 1982. Seasonal patterns of dormancy in weed seeds. Pages
123-145 in A. A. Khan, ed. The Physiology and Biochemistry of Seed
Development, Dormancy and Germination. Amsterdam: ElsevierBio-
medical Press.
Kropf, M. J. and H. H. Van Laar. 1993. Modeling crop-weed interactions.
Wallingford: CAB International. pp. 105-133
Lafond, G. P. and R. J. Baker. 1986. Effects of genotype and seed size on
speed of emergence and seedling vigor in nine spring wheat cultivars.
Crop Sci. 26:341-346.
Lueshen, W. E. and R. N. Andersen. 1980. Longevity of velvetleaf (Abutilon
theophrasu") seeds in soil under agricultural practices. Weed Sci. 28:
341-346.
Mapes, G., G. W. Rothwell, and M. T. Haworth. 1989. Evolution of seed
dormancy. Nature 337:645-646.
Mester, T. C. and D. D. Buhler. 1991. Effects of temperature, seed depth,
and cyanazine on giant foxtail (Setariafaberi) and velvetleaf (Abutilon
theophrasti)seedling development. Weed Sci. 39:204-209.
The author has requested enhancement of the downloaded file. All in-text references underlined in blue are linked to publications on ResearchGate.
Milberg, P., L. Andersson, C. Elfverson, and S. Regner. 1996. Germination
characteristicsof seeds differing in mass. Seed Sci. Res. 6:191-197.
Mitich, L. W. 1991. Field bindweed. Weed Technol. 5:913-915.
Mohler, C. L. 1993. A model of the effects of tillage on emergence of weed
seedlings. Ecol. Appl. 3:53-73.
Mohler, C. L. and A. E. Galford. 1997. Weed seedling emergence and
survival: separating the effects of seed position and soil modification
by tillage. Weed Res. 37:147-155.
Pareja, M. R. and D. W. Staniforth. 1985. Seed-soil characteristicsin re-
lation to weed seed germination. Weed Sci. 33:190-195.
Pons, T. L. 1991. Induction of dark dormancy in seeds: its importance for
the seed bank in the soil. Funct. Ecol. 5:669-675.
Raymond, P., A. Al-Ani, and A. Pradet. 1985. ATP production by respi-
ration and fermentation, and energy charge during anaerobiosis and
aerobiosis in twelve fatty and starchy germinating seeds. Plant Physiol.
79:879-884.
Roberts, E. H. and S. Totterdell. 1981. Seed dormancy in Rumex species
in response to environmental factors. Plant Cell Environ. 4:97-106.
Roberts, H. A. and M. E. Potter. 1985. Emergence patterns of weed seed-
ling in relation to cultivation and rainfall. Weed Res. 30:377-382.
Roberts, H. A. and M. E. Ricketts. 1979. Quantitative relationshipbetween
the weed flora after cultivation and the seed population in the soil.
Weed Res. 19:269-275.
Stoller, E. W. and L. M. Wax. 1973. Periodicity of germination and emer-
gence of some annual weeds. Weed Sci. 21:574-580.
Taylor, A. G. and C. W. Ten Broeck. 1980. Seedling emergence forces of
vegetable crops. J. Hortic. Sci. 23:367-369.
Terpstra, R. 1995. Dormancy of seeds of shepherd's purse in alternating
wet and dry, compressed aggregated soil: a laboratory experiment. J.
Appl. Ecol. 32:434-444.
Thompson, K. 1987. Seeds and seedbank. New Phytol. 106(Suppl.):23-
34.
Thompson, K., S. R. Band, and J. G. Hodgson. 1993. Seed size and shape
predict persistence in soil. Funct. Ecol. 7:236-241.
Vleeshouwers, L. M. 1997. Modelling the effect of temperature, soil pen-
etration resistence, burial depth and seed weight on pre-emergence
growth of weeds. Ann. Bot. 79:553-563.
Weaver, K. N. 1980. Field emergence of calabrese and onion seedlings in
response to compaction treatments on the soil surfaceor at seed depth.
J. Hortic. Sci. 55:325-332.
Webb, D. M., C. W. Smith, and J. Schulz-Scaeffer. 1987. Amaranth seed-
ling emergence as affected by seeding depth and temperature on a
thermogradient plate. Agron. J. 79:23-26.
Wesson, G. and P. E Wareing. 1969. The induction of light sensitivity in
weed seeds by burial. J. Exp. Bot. 20:414-425.
Wiese, A. F and L. K. Binning. 1987. Calculating the threshold temper-
ature of development of various weeds. Weed Sci. 35:177-179.
Wiese, A. F and R. C. Davis. 1967. Weed emergence from two soils at
various moistures, temperatures,and depths. Weeds 15:118-121.
Zhang, J., A. S. Hamill, 1. 0. Gardiner, and S. W. Weaver. 1998. Depen-
dence of weed flora on the active soil seedbank. Weed Res. 38:143-
152.
ReceivedFebruary4, 2000, and approvedMarch 13, 2001.
Benvenutiet al.: Buriedweed seedlingemergence * 535