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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 .
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WeedScience,49:528-535. 2001

Quantitative analysis of emergence of seedlings from


buried weed seeds with increasing soil depth

Stefano Benvenuti Trialswere carriedout to investigatethe effectsof seed burialdepth on seedling


Corresponding author. Dipartimento di Agronomia emergencerateof 20 weed species.Markeddepth-mediatedvariationin emergence
e Gestione dell'Agroecosistema,Universita di Pisa, abilityof the differentspecieswas observed,togetherwith a generalpatternof de-
Via S.Michele degli Scalzi 2, 56124 Pisa, Italy; creasingemergencewith increasingsoil depth.At 10 cm, only johnsongrass, velvet-
sbenve@agr.unipi.it leaf,catchweedbedstraw,and cutleafgeraniumemerged,albeitonly in limitednum-
bers. Speciesmost severelyinhibitedby burialdepthwerebuckhornplantain,large
Mario Macchia crabgrass,common purslane,chickweed,and corn spurry,none of which emerged
Sergio Miele from beyond 6 cm. In all species,depth-mediatedinhibitionwas found to be sig-
Dipartimento di Agronomia e Gestione moidal (polynomialregression).In addition,the numberof seedlingsand rate of
dell'Agroecosistema,Universita di Pisa, Via seedlingemergencedecreasedwhen depth of burialincreased.The depth at which
S.Michele degli Scalzi 2, 56124 Pisa, Italy 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.

Knowledge of weed biology has aroused increasing inter- Germination is also known to be inhibited by soil depth
est in recent years in the context of weed management strat- (Holm 1972; Stoller and Wax 1973). The biological reason
egies (Bhowmik 1997). Such knowledge is essential to max- for depth inhibition has not yet been fully clarified. Previous
imize the effectiveness of agronomic practices (Forcellaet al. studies suggest this may not be due merely to lack of light
1993) and to simulate weed dynamics. The greater insight (Benvenuti 1995), but also to decreasingthermal fluctuation
into weed dynamics gained through this approach has with increasing burial depth, as thermal fluctuation consti-
opened up new horizons in crop protection, allowing ratio- tutes a known germination trigger (Roberts and Totterdell
nal utilization of chemical control. The overall aim is to 1981). An alternative or possibly complementary explana-
apply herbicides exclusively, and proportionately, only in tion is based on gas diffusion, which is inversely correlated
cases of genuine risk of weed-crop competition. with burial depth. Reduced gas exchange is believed to be
Weed dynamics simulations have considered a number of capable of inducing secondary dormancy (Benvenuti and
patterns of the weed life cycle, including agronomic soil Macchia 1995). Presence of CO2 deriving from soil biolog-
disturbance (Ball 1992; Mohler 1993), response to ecolog- ical activity is probably also involved in this complex mech-
ical and climatic factors (Alm et al. 1993; Forcella 1993, anism (Karssen 1982). It is therefore of prime importance
1998), and even subsequent weed growth within crops to acquire knowledge on weed seed distribution in soil
(Kropf and Van Laar 1993). Crucial to successful simulation (Cousens and Moss 1990; Grundy et al. 1996) and on the
is knowledge of the extent of the soil seed bank (Forcella emergence ability of various weed species (Alm et al. 1993).
1992; Roberts and Ricketts 1979; Zhang et al. 1998), an- Emergence from different soil depths has been found to be
nual seed production (Benvenuti et al. 1994; Cardina and proportional to seed energy reserves (Lafond and Baker
Norquay 1997), and aspects of buried seed ecology (Pons 1986). It is also influenced by the typology of such reserves,
1991) that can influence seed longevity (Lueshen and An- depending on the amount of oxygen required for their uti-
dersen 1980). Ecological factors have been shown to play a lization (Raymond et al. 1985). Energy reservesare vital for
major role, not only by influencing secondary dormancy seedling growth prior to emergence, because given the ab-
(Baskin and Baskin 1985) but also by either inducing or sence of light, growth takes place completely autotrophically.
inhibiting germination. Thus, it is well known that light Furthermore, independent of the physiological aspects char-
(Ballare et al. 1992), temperature (Benvenuti and Macchia acterizing this agronomically important stage, weed species
1993), soil water content (Roberts and Potter 1985), and may differ in their ability to emerge from the various soil
degree of soil compaction (Parejaand Staniforth 1985) rep- layers.
resent the main factors limiting buried seed germination. Accurate knowledge of these parameters exerts a marked

528 * Weed Science 49, July-August 2001


influenceon the degreeof precisionof weed dynamicsim- Seedson the soil surfacewere coveredwith moistenedfilter
ulations.It consequentlyaffectssuccessfulagronomicappli- paper1at first (12 h) to stimulateimbibition.
cation of weed dynamicsmodelingand, thus, the possibility
of setting up rationalweed controlstrategies. Buried Seed Incubation
The purposeof this workwas (1) to performquantitative
analysisof the seedlingemergencecharacteristicsof 20 dif- Seeded pots were placed in climate-controlledcabinets
ferentweed speciesas a function of depth of burialand (2) presetto alternatingtemperaturesof 25/30 C for predom-
to investigatepossiblecorrelationsbetweenemergenceabil- inantly summer weeds (prostrateknotweed, redroot pig-
ity and seed size. weed, jimsonweed,Canadathistle,largecrabgrass,johnson-
grass, barnyardgrass, common lambsquarters,black night-
shade, velvetleaf,field bindweed, and common purslane)
Materials and Methods and 15/20 C for predominantlyspring weeds (buckhorn
Trialswere carriedout at the Seed Researchand Testing plantain,blackgrass,wild mustard,hairybittercress,catch-
Station(InternationalSeedTestingAssociationapproved)of weed bedstraw,cutleafgeranium,common chickweed,and
the AgronomyDepartmentof Pisa University,Italy.Seeds corn spurry).The same 12-h dark/lightphotoperiodwas
of 20 weed species-prostrate knotweed (Polygonumla- maintainedfor both temperatures.Light intensityof 100
phathifoliumL. POLLA),redrootpigweed (Amaranthus re- Lmol m-2 s-1 was produced by cool fluorescent tubes2 and
troflexusL. AMARE),buckhornplantain,jimsonweed(Da- was measuredwith a spectroradiometer.3 During incuba-
tura stramoniumL. DATST), Canadathistle [Cirsiumar- tion, pots were moistenedby subirrigation.
vense(L.) Scop. CIRAR], large crabgrass,blackgrass(Alo-
pecurus myosuroides Huds. ALOMY), johnsongrass, Seedling Emergence
barnyardgrass [Echinochloacrus-galli(L.) Beauv.ECHCG],
wild mustard[Brassicakaber(DC.) L.C. WheelerSINAR], Emergedseedlingswere counted daily at cotyledonap-
common lambsquarters(Chenopodium albumL. CHEAL), pearanceand removed.Seedlingcounts were stopped3 to
black nightshade(SolanumnigrumL. SOLNI), velvetleaf 5 d afterno moreemergencewas recorded.Meanemergence
(Abutilontheophrasti),hairybittercress(CardaminehirsutaL. time (MET) was calculatedas
CARHI), catchweedbedstraw,cutleaf geranium,common MET = l(n x g)lN,
purslane,field bindweed(Convolvolus arvensisL. CONAR),
common chickweed,and corn spurry-were collectedat full where n is the numberof seedlingsemergingper day,g is
ripeningin spring(une 15) and summer(August25) 1998 the number of days needed for emergence,and N is the
near Pisa, Italy.Seeds were cleanedimmediatelyafter har- total numberof emergedseeds.Foreachseedingdepth,per-
vesting and placed in hermeticallysealedscrew-topplastic centageof soil depth inhibitionwas calculatedfor eachspe-
jars.Seedswerestoredin darknessat room temperature(20 cies as a function of unburiedseed germination(0% inhi-
C) and relativehumidity < 10%, determinedaccordingto bition).
ISTA methods (1999) based on the loss in weight when
seedsaredrieduntil use in September1999. Beforeseeding, Ungerminated Seed Recovery and Germination
18 weed specieswerestratifiedat 4 C for 2 wk to overcome Test
any dormancy.An exceptionto this procedurewas adopted
for velvetleafand field bindweed(Horowitzand Taylorson After emergencetests, soil was removedfrom pots and
1985; Mitich 1991, respectively),whichwereburied(in jute washed to determinethe fate of ungerminatedseeds (dor-
bags)in the field 30 cm deep for 1 yr priorto use (Septem- mant and/or germinatedwithout emergence).A fine metal
ber 1998-September1999) to overcomecharacteristic seed sieve (400 gm) was used for seed recovery.Suicidegermi-
hardness,according to a previouslydescribedprocedure nation (not followedby seedlingemergence)was calculated
(Cardinaand Sparrow1997). as the differencebetweentotal seedsand the sum of recov-
ered seedsand emergedseedlings.Fifty ungerminatedseeds
Seeding by each pot from the deep burialof 12 cm were imbibed
with 3 ml of distilledwateron a single sheet of filterpaper
Fifty seeds of each of the 20 weed specieswere sown in placedin 6-cm petri dishes.Germinationtestswere carried
plasticpots (15 by 15 by 25 cm) filledwith a silt-loamsoil out in the same photothermicconditionsas the emergence
(typic xerofluvent soil; 23% sand, 52% silt, 25% clay). Soil tests.
was obtainedby excavationsfrom a depth of over 1 m in
orderto avoid the presenceof preexistingseeds capableof
Seed Weight
falsifyingthe real experimentaldata of emergencerates.In
spite of obtainingsoil froma deep soil layer,the relativesoil Seed weight was determinedby weighing 1,000 seeds
characteristicswere similar to a normal topsoil (pH 6.8, chosen randomlyaccordingto ISTA rules for seed testing
0.08% total N, 21 mg kg-1 availableP205, 265 mg kg-l (ISTA 1999).
exchangeableK20) and no particularproblembasedon aer-
obic or anaerobicconditions was found. Pots were filled Calculation of Depth of 50% Emergence
gravimetricallywith the soil and packed with a uniform Inhibition
strength to avoid differential resistance to seedling emer-
gence. Seeding depths were 0, 2, 4, 6, 8, 10, or 12 cm. The Polynomial regressions were calculated that showed the
study was a completely randomized design with three rep- best fit of the biological response of weed seed emergence-
licates for each seeding depth and the study was repeated. 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 essary.Angular values were subjected to analysis of variance
soil depths capable of reducing emergence to 50% of soil (ANOVA) using the Student-Newman-Keuls test (P <
surface germination. Depth values were plotted with the 0.05) for means separation. Inhibition data of each species
corresponding 1,000-seed weight and fitted with a quadratic tested were fitted by the corresponding polynomial regres-
polynomial regression. 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%
Statistical Analysis by using a modified "x-intercept"method (Wiese and Bin-
The two experiments did not differ in seed response and ning 1987). The intercept between the polynomial regres-
data were pooled. After testing for homogeneity of variance, sion and the translated x-axis on the selected y for 50%
arcsine transformation of emergence percentages was nec- 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- shows that prostrate knotweed, redroot pigweed, and jim-
ware (CoStat4) was used. sonweed had emergence less than 10% at a burial depth of
8 cm, whereas buckhorn plantain showed complete inhibi-
Results and Discussion tion at this depth. Large crabgrasswas unable to emerge
from a depth greater than 6 cm, whereas johnsongrass did,
Figure 1 shows the emergence of the 20 weed species albeit at a minimum percentage (roughly 5%), from as deep
studied as a function of depth of burial. Emergence de- as 10 cm. However, for the majority of species such as Can-
creased with increasing burial depth, with 12 cm represent- ada thistle, blackgrass,barnyardgrass,common lambsquart-
ing the limit of total inhibition at which no species was able ers, black nightshade, and wild mustard, the depth limit was
to emerge. For each of the species examined, depth inhibi- considerably less then 10 cm, and emergence was very low
tion was found to be highly significant (P < 0.01) by fitting (ranging between 5 and 10%) even at a burial depth of 8
the relevant data by polynomial regression. cm. The greatest ability to maintain emergence with increas-
Analysis of the outcome for individual species (Table 1) 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, shaw 1992; Mester and Buhler 1991; Vleeshouwers 1997;
which were generally able to germinate and reach the soil Webb et al. 1987).
surface even from 10 cm (5 to 15%). In contrast, hairy In our study, burial depth also exerted marked influence
bittercress, common purslane, common chickweed, and on MET in each of the species tested (Table 1). Each 2 cm
corn spurry showed marked inability to emerge from deep increase in distance below the soil surface resulted in a sig-
burial and a limit of about 6 cm. nificant (P < 0.05) increase in emergence time. This delay
In all species tested, emergence decreased slightly with in emergence from deeper soil levels means that deeply bur-
shallow burial (2 cm) and then decreased exponentially at ied seeds have a disadvantage in competition with crops.
greater depths. However, some authors (Chancellor 1964; Use of equations to representinhibition induced by burial
Mohler and Galford 1997; Wiese and Davis 1967), who depth (expressedas a percentage compared to unburied seed
performed similar experiments on analogous weed species, germination) allowed identification of the burial depth at
reported a favorable effect of slight burial (roughly 0.5 to 1 which emergence was halved. This depth varied depending
cm) on emergence. Such an effect was not detected in the on the species examined, ranging from 3.6 cm in common
present study. This could have been because of water stress purslane and common chickweed to 7 cm with velvetleaf
near the soil surface in the cited studies, a phenomenon that and catchweed bedstraw. Along this gradient, Canada this-
was avoided in the present test by covering the soil surface tle, black nightshade, and barnyardgrassshowed depth-me-
with filter paper. Additionally, in our study, the first depth diated emergence (5.3 cm) approaching mean 50% emer-
below the surface layer studied was 2 cm, a depth that did gence values of the 20 species tested (mean 5.1 cm). The
not include the zone in which variable responses were de- level of emergence we found was slightly greaterthan values
scribed in the cited studies. detected in field conditions by other authors (reviewed by
Other factors leading to difficulty in comparing results Mohler 1993). This discrepancy could be due to the greater
obtained in this study with those reported by other authors physical constriction in a field with undisturbed soil than in
include possible differences in dormancy characteristicsof our dry-sieved experimental soil. It has been shown that soil
the various species used, the particular temperature and hu- compaction impairs the ability of seeds to reach the soil
midity conditions, and above all, the different soil charac- surface during preemergence seedling growth (Hegarty and
teristics. Soil particle size has been shown to influence soil Royale 1978). Furthermore,soil compaction can act directly
physical characteristics,which can interferewith buried seed on seeds by limiting germination (Pereja and Staniforth
germination and emergence (Cussans et al. 1996). Despite 1985) or even by inducing dormancy (Terpstra1995). How-
these general differences, there is overall agreement that ever in our study, the effects of soil compaction were not
emergence inhibition increases proportionately with depth examined. A relative comparison was made of depth-medi-
of seed burial in soil (Benvenuti and Macchia 1997; Black- 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


that involve or simulate seed bank dynamics. Only by
C) 4-1 .- achievingprecisionin seed dynamicsimulationwill it be-
come possible to make weed control optimally effective
7My 0 while minimizingunjustifiedherbicideapplication.This will
favor harmonizationof ecologicalconcernswith the eco-
a 30
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C 08 nomic advantagesexpectedfrom modernagriculture.
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Sources of Materials
eq ( y=- 0.036 x2he0.639 x+ 4.12
1 Filter paper, Watchman No. 1. Fisher Scientific, Pittsburgh,PA
o o R 0.75 15219.
FIUE 3. Rersinbtenbra0et eesr ordc mrec 2 Fluorescent lamp, Philips TLF 20W/33, Eindhoven,
The
- 4-- 0
Netherlands.
n 0 1 2 3 4 5 6 7 8 9 10 11 12 3 Spectroradiometermodel 1800. Licor, Inc., Lincoln, NE.
4 CoHort Software, Minneapolis, MN 55419.
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. Literature Cited
The equation (significant for P > 0.05) and the corresponding R2 value
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Benvenutiet al.: Buriedweed seedlingemergence * 535


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