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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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Stellariamedia(L. Accurate knowledge of these parameters exerts a marked 528 * Weed Science 49. only in tion is based on gas diffusion. Roberts and Ricketts 1979. Zhang et al. Italy the numberof emergedseedlingswas halvedvariedby speciesand rangedfrom3. Germination is also known to be inhibited by soil depth est in recent years in the context of weed management strat. depending on the amount of oxygen required for their uti- dersen 1980).chickweed.The depth at which S. Grundy et al. not only by influencing secondary dormancy seedling growth prior to emergence. Cardina and proportional to seed energy reserves (Lafond and Baker Norquay 1997). only johnsongrass. The greater insight (Benvenuti 1995). Key words: Seed germination. temperature (Benvenuti and Macchia acterizing this agronomically important stage. Via seedlingemergencedecreasedwhen depth of burialincreased.Universita di Pisa.seedlingemergence.it leaf.Excessiveburialdepth generallyinduceddormancy(in roughly85% of cases)ratherthan suicidegermination.SORHA. DIGSA. Crucial to successful simulation (Cousens and Moss 1990.corn spurry.common purslane. Presence of CO2 deriving from soil biolog- disturbance (Ball 1992.6 cm for common purslaneand chickweedto 7 cm for velvetleafand catchweed bedstraw.WeedScience. 56124 Pisa. Nomenclature: Buckhornplantain.and corn spurry. An alternative or possibly complementary explana- apply herbicides exclusively. allowing ratio. Ecological factors have been shown to play a lization (Raymond et al.) Scop. and may differ in their ability to emerge from the various soil degree of soil compaction (Parejaand Staniforth 1985) rep.none of which emerged Sergio Miele from beyond 6 cm.A close inverserelation(second-degreeequa- tion) betweenseed unit weight and depth-mediatedinhibitionwas observed. Forcella 1993.togetherwith a generalpatternof de- Via S. Mohler 1993).johnsongrass. including agronomic soil Macchia 1995).common chickweed. anism (Karssen 1982).Michele degli Scalzi 2.cutleafgeranium. 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.PlantagolanceolataL. SPRAR.The physiologicalinvolvementof depth inhibitionin seed bankecologyis discussed. 1993. soil water content (Roberts and Potter 1985). It is therefore of prime importance 1998). 1994. Energy reservesare vital for major role.largecrabgrass. Previous imize the effectiveness of agronomic practices (Forcellaet al.depth-mediatedinhibitionwas found to be sig- Dipartimento di Agronomia e Gestione moidal (polynomialregression). 1992). independent of the physiological aspects char- (Ballare et al. resent the main factors limiting buried seed germination. GaliumaparineL. Knowledge of weed biology has aroused increasing inter. GALAP.49:528-535. as thermal fluctuation consti- opened up new horizons in crop protection. 1992. Thus.In addition. Dipartimento di Agronomia emergencerateof 20 weed species.) Vill.Abutilontheophrasti MedicusABUTH. STEME.common purslane.) Pers.PortulacaoleraceaL. 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. Such knowledge is essential to max.Markeddepth-mediatedvariationin emergence e Gestione dell'Agroecosistema.At 10 cm.velvetleaf. an. because given the ab- (Baskin and Baskin 1985) but also by either inducing or sence of light. inhibiting germination. 1993). 1991) that can influence seed longevity (Lueshen and An. growth takes place completely autotrophically. The overall aim is to 1981). it is well known that light Furthermore.Universita di Pisa. creasingemergencewith increasingsoil depth. Sorgumhalepense(L.catchweedbedstraw. layers. In all species. velvet- sbenve@agr.albeitonly in limitednum- bers.Michele degli Scalzi 2.Geraniumdissectum L. but also to decreasingthermal fluctuation into weed dynamics gained through this approach has with increasing burial depth. and proportionately. 1998). PLALA. The biological reason egies (Bhowmik 1997). (Holm 1972.Spergula arvensL. 1996) and on the is knowledge of the extent of the soil seed bank (Forcella emergence ability of various weed species (Alm et al.soil seed bank.catchweedbed- straw.large Mario Macchia crabgrass. abilityof the differentspecieswas observed. weed species 1993). response to ecolog. and even subsequent weed growth within crops to acquire knowledge on weed seed distribution in soil (Kropf and Van Laar 1993). Speciesmost severelyinhibitedby burialdepthwerebuckhornplantain.burialdepth. and aspects of buried seed ecology (Pons 1986). Stoller and Wax 1973).the numberof seedlingsand rate of dell'Agroecosistema. with burial depth. It is also influenced by the typology of such reserves. GERDI. studies suggest this may not be due merely to lack of light 1993) and to simulate weed dynamics.Digitariasanguinalis(L. for depth inhibition has not yet been fully clarified.and cutleafgeraniumemerged. POROL. Italy. which is inversely correlated cases of genuine risk of weed-crop competition. July-August 2001 . ical activity is probably also involved in this complex mech- ical and climatic factors (Alm et al. Emergence from different soil depths has been found to be nual seed production (Benvenuti et al. 56124 Pisa. tutes a known germination trigger (Roberts and Totterdell nal utilization of chemical control.unipi. 1985).

4. 8. 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. 265 mg kg-l (ISTA 1999). 10.Suicidegermi- hardness.common chickweed. Buried Seed Incubation The purposeof this workwas (1) to performquantitative analysisof the seedlingemergencecharacteristicsof 20 dif. 52% silt.A fine metal ber 1998-September1999) to overcomecharacteristic seed sieve (400 gm) was used for seed recovery.common MET = l(n x g)lN. Mitich 1991. and common purslane) Materials and Methods and 15/20 C for predominantlyspring weeds (buckhorn Trialswere carriedout at the Seed Researchand Testing plantain. thus. maintainedfor both temperatures. pearanceand removed.according to a previouslydescribedprocedure nation (not followedby seedlingemergence)was calculated (Cardinaand Sparrow1997). The Polynomial regressions were calculated that showed the study was a completely randomized design with three rep. Italy. Canadathistle [Cirsiumar.influenceon the degreeof precisionof weed dynamicsim.The same 12-h dark/lightphotoperiodwas of 20 weed species-prostrate knotweed (Polygonumla. total numberof emergedseeds.field bindweed. CHEAL).Germinationtestswere carried plasticpots (15 by 15 by 25 cm) filledwith a silt-loamsoil out in the same photothermicconditionsas the emergence (typic xerofluvent soil.) Beauv. paper1at first (12 h) to stimulateimbibition.In spite of obtainingsoil froma deep soil layer. DATST). CONAR).Light intensityof 100 phathifoliumL. as the differencebetweentotal seedsand the sum of recov- ered seedsand emergedseedlings.Canadathistle. SOLNI).) Scop.blackgrass. mant and/or germinatedwithout emergence). ISTA methods (1999) based on the loss in weight when seedsaredrieduntil use in September1999. 21 mg kg-1 availableP205. weed. inhibition at increasing soil depths.Seedlingcounts were stopped3 to black nightshade(SolanumnigrumL. 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. common chickweed.buckhornplantain. determinedaccordingto bition).field bindweed(Convolvolus arvensisL. Seedling Emergence barnyardgrass [Echinochloacrus-galli(L. Ungerminated Seed Recovery and Germination 18 weed specieswerestratifiedat 4 C for 2 wk to overcome Test any dormancy. chosen randomlyaccordingto ISTA rules for seed testing 0. 2. or 12 cm. time (MET) was calculatedas CARHI). pots were moistenedby subirrigation.C.) L. 23% sand. Beforeseeding.black night- shade. AMARE).Seeds corn spurry).jimsonweed(Da.: Buriedweed seedlingemergence * 529 . large crabgrass.An exceptionto this procedurewas adopted for velvetleafand field bindweed(Horowitzand Taylorson After emergencetests.whichwereburied(in jute washed to determinethe fate of ungerminatedseeds (dor- bags)in the field 30 cm deep for 1 yr priorto use (Septem.hairybittercress(CardaminehirsutaL. tion. johnsongrass.blackgrass(Alo- pecurus myosuroides Huds. respectively). the possibility of setting up rationalweed controlstrategies. Emergedseedlingswere counted daily at cotyledonap- common lambsquarters(Chenopodium albumL.cutleafgeranium.ECHCG]. purslane.johnson- grass.08% total N. WheelerSINAR]. Seedson the soil surfacewere coveredwith moistenedfilter ulations. soil was removedfrom pots and 1985.and corn spurry-were collectedat full where n is the numberof seedlingsemergingper day.hairybittercress.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.Italy. velvetleaf. was measuredwith a spectroradiometer.8.catch- Station(InternationalSeedTestingAssociationapproved)of weed bedstraw. velvetleaf 5 d afterno moreemergencewas recorded. CIRAR].3 During incuba- tura stramoniumL.000 seeds characteristicswere similar to a normal topsoil (pH 6.redrootpigweed (Amaranthus re. barnyardgrass. Soil tests. These polynomial equa- Benvenutiet al. exchangeableK20) and no particularproblembasedon aer- obic or anaerobicconditions was found. ALOMY). cation of weed dynamicsmodelingand. Lmol m-2 s-1 was produced by cool fluorescent tubes2 and troflexusL. vense(L. jimsonweed. best fit of the biological response of weed seed emergence- licates for each seeding depth and the study was repeated.Foreachseedingdepth.Seedswerestoredin darknessat room temperature(20 cies as a function of unburiedseed germination(0% inhi- C) and relativehumidity < 10%. was obtainedby excavationsfrom a depth of over 1 m in orderto avoid the presenceof preexistingseeds capableof Seed Weight falsifyingthe real experimentaldata of emergencerates. inantly summer weeds (prostrateknotweed. common lambsquarters.the relativesoil Seed weight was determinedby weighing 1.cutleaf geranium. Seeding depths were 0.g is ripeningin spring(une 15) and summer(August25) 1998 the number of days needed for emergence.wild mustard.and N is the near Pisa.and the AgronomyDepartmentof Pisa University. 25% clay).per- vesting and placed in hermeticallysealedscrew-topplastic centageof soil depth inhibitionwas calculatedfor eachspe- jars.Meanemergence (Abutilontheophrasti).Seeds were cleanedimmediatelyafter har.largecrabgrass.It consequentlyaffectssuccessfulagronomicappli. POLLA). 6. catchweedbedstraw. redroot pig- ity and seed size. wild mustard[Brassicakaber(DC.

July-August 2001 .0 j2 09x+03 100 so 3.9 100 R20. Depth values were plotted with the 0.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.84+1 13 ? i100 ? uz 3o / 80 C:so.61 x-0. tions of soil depth inhibition activity were used to identify essary. sion and the translated x-axis on the selected y for 50% arcsine transformation of emergence percentages was nec.98 :r.01) and the corresponding R2 values are reported.024x-o027 1 +5.20-6 . 100 . a + r20Xta e-N20da / 00 eoL g lCda ethistle Large crabgrass e a C LI~~~~~~R-09 R -0. 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=-0097x -1 1.98 9 80 so~~~~~~~~~~~~~~~~~~~~~~~~8 Prostrate knotweed Redroot pigweed 40 40 y--0.000-seed weight and fitted with a quadratic tested were fitted by the corresponding polynomial regres- polynomial regression. depth inhibition shows the relative soil depth inhibition for 530 * Weed Science 49. _=016Y25I00 2=ooe -Fi~ x7.34x1 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. . [0 1 k 05260 6x+5.9x-03 10 6. y-0. sion that adequately described the biological response of weed seed germination and emergence. y--0.05 according to the Student-Newman-Keuls test.1~~~~~~~~~~~~~~.038x4-1 31i &1875x-2.121 xi .57x3-1 S x . Means followed by the same letter do not differ at P < 0. y--=Ol 73x+2.28x-3 4 - _.92xA2.l1 4x-2. After testing for homogeneity of variance. Inhibition data of each species corresponding 1. Arrows indicate the values of 50% depth inhibition. y--0.05) for means separation. The equations of inhibitions (significant for P < 0.98 > R2=0.45x-4. The intercept between the polynomial regres- data were pooled.85 b so8 60 1 i/ t z Jimsonweed ~~~~Buckhorn plantain 100 . 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).ef 0 '|~~~~~~~~~~~~~~~~~~~~~~-.3 .

was shown by velvetleaf.01) by fitting (ranging between 5 and 10%) even at a burial depth of 8 the relevant data by polynomial regression. However. as 10 cm.4 . Benvenuti et al. depth inhibi. considerably less then 10 cm.87x-i-08 .C66 100 ff 80~~~ Common lambsquarters| Black nightshade R2-0. and emergence was very low tion was found to be highly significant (P < 0. commercial soft.56x-3. the depth limit was to emerge.98 > so0 a so e01 a x.: Buried weed seedling emergence * 531 .~~~ Common7. Continued.30x2. For each of the species examined. from as deep studied as a function of depth of burial. ada thistle. The greatest ability to maintain emergence with increas- Analysis of the outcome for individual species (Table 1) ing depth. whereas johnsongrass did. sonweed had emergence less than 10% at a burial depth of 8 cm. For each statistical analysis. although at low levels. redroot pigweed. y--0. 7-1 1~. < d d | 1 /8 X e e 2 100 _y=-0. whereas buckhorn plantain showed complete inhibi- Results and Discussion tion at this depth.barnyardgrass. cm. 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. 1 FIGURE_1. 100 -Y-0.. Figure 1 shows the emergence of the 20 weed species albeit at a minimum percentage (roughly 5%). shows that prostrate knotweed. and wild mustard. with 12 cm represent.common lambsquart- ing the limit of total inhibition at which no species was able ers. rlambqurtr so~~~~~~~~~~~~~~~~~~~5 a SX a 150 -a 20L 50 .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. Commonept purian bedn Fieldhbinwee each species.97 Rf -0. black nightshade.170X+3. Large crabgrasswas unable to emerge from a depth greater than 6 cm.58x-3.gBx-5. and jim- ware (CoStat4) was used. Emergence de. =OeOt 1fh i .152A2.149xt-2. for the majority of species such as Can- creased with increasing burial depth. blackgrass. Continued.

In contrast.2 (3. Additionally.5 (3. Such an effect was not detected in the on the species examined. Alopecurusmyosuroides.9) 17.3 cm) approaching mean 50% emer- below the surface layer studied was 2 cm.velvetleaf. cutleaf geranium.0 (1.1) 7. and barnyardgrassshowed depth-me- with filter paper.1 (1. 1996).2) Cutleaf geranium 7.4 (1.7) 7.5) 11.9) 23.3) 7.TABLE 1.7 (2. there is overall agreement that ever in our study.johnsongrass. germination) allowed identification of the burial depth at reported a favorable effect of slight burial (roughly 0.4 (1. Spergula arvensis. Portulaca oleracea. compaction impairs the ability of seeds to reach the soil midity conditions.2 (1.0 (0.8 (0.5) 9.8 (0.6 (0.1) Wild mustard 3.8) Corn spurry 6.8) 10.5) 5. 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.4 (1.4 (0. and on MET in each of the species tested (Table 1). Use of equations to representinhibition induced by burial Mohler and Galford 1997.1 (1.3) 13.5 (1.0) 23.field bindweed.3) 20. 1987). burial depth also exerted marked influence bittercress. - a Barnyardgrass.8 (1.3 (1.8) 10.0) 21. hairy In our study.4) 12.4) 12.5) 16.9) Redroot pigweed 4.9) 11. It has been shown that soil the various species used. catchweed bedstraw. Canadathistle.5) 7.7 (2. Cirsium arvense.6) 7.2 (1.3) 8. Soil particle size has been shown to influence soil Royale 1978).0) 10.2 (2.) Meanemergencetime (seedingdepth.7 (0. Stellaria media.catchweed bedstraw.prostrateknotweed. Amaranthusretroflexus-. in our study.8 (0.5) 7.2 (1. 532 * Weed Science 49.0) 9.9) Hairy bittercress 4.9) 22.5) 13. (Means are followed by the relative standard errors. Chenopodiumalbum.5 to 1 which emergence was halved.1 (0.7 (1. However.7 (2. This delay In all species tested.4 (0. Canada this- was avoided in the present test by covering the soil surface tle.1) 5.7) 17. Plantago lanceolata.3 (1.2) 19.9) 16.8 (0.2) 19.6 (0.5 (0. some authors (Chancellor 1964. a phenomenon that and catchweed bedstraw.8 (0.5 (3.7) Barnyardgrass 6.large crabgrass. blackgrass.3) 9.2 (2.1) . nificant (P < 0. black nightshade. surface even from 10 cm (5 to 15%).9 (1. common purslane. Effect of seeding depth on mean emergence time of the 20 tested weed species.5 (2.corn spurry. This depth varied depending cm) on emergence.6 (0.05) increase in emergence time.5 (0.2 (3. Abutilon theophrasti. 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.1 (2.0 (1.4 (0.6) 13.7) Common chickweed 5. Polygonumlaphathifolium.9 (0.3) 7.3 (0.6) 15.8 (1.9) 12. A relative comparison was made of depth-medi- of seed burial in soil (Benvenuti and Macchia 1997. The not include the zone in which variable responses were de.4) 10.3) 8.8) 9. ranging from 3.5 (1. Despite 1985) or even by inducing dormancy (Terpstra1995).1) Common lambsquarters 6.5 (0. ated emergence among different weed species under similar.3) 8. cutleaf geranium.6 (3.buckhorn plantain.1 (0. Brassicakaber. Geranium dissectum.7) Catchweed bedstraw 6.which can interferewith buried seed on seeds by limiting germination (Pereja and Staniforth germination and emergence (Cussans et al. Mester and Buhler 1991.6) 6.2 (1.6) 7. How- these general differences.3) 9.0) 16.5) 19. Along this gradient.2 (0.8 (0. the different soil charac. common purslane.0) Black nightshade 7. the particular temperature and hu. black nightshade.2 (0. shaw 1992.jimsonweed.2 (1. a depth that did gence values of the 20 species tested (mean 5.8) 12.3 (0.5 (0. commonchickweed.9 (0.1 (2.1 (3.5 (0.2) 13.2) 10. commonlambsquarters.9) 11.2 (1.7 (0.5 (0. Furthermore.1) 12.7 (0.6 cm in common present study.3 (0.4) 20. 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.9) 12. the first depth diated emergence (5.8) 12.6 (1.7) 10.2) 9.4) 6. Sorghumhalepense. Cardaminehirsuta. surface during preemergence seedling growth (Hegarty and teristics. Solanum nigrum.2 (2.8 (1.1 (0.1 (1. detected in field conditions by other authors (reviewed by Other factors leading to difficulty in comparing results Mohler 1993). Wiese and Davis 1967).4) 12.redrootpigweed.1 (1.7 (0. Galium aparine.1 cm).3 (0.5 (0.8) 8.2) 11.9) 13.4) 6. greater depths.1) Jimsonweed 7. who depth (expressedas a percentage compared to unburied seed performed similar experiments on analogous weed species.6) 7.4) Field bindweed 6.6 (3.4 (1.7) 8.1 (1.4) 11. b Missing values indicate no seedling emergence.1 (2.3 (0. July-August 2001 . Convolvolusarvensis.6) 19.2) Velvetleaf 5.0 (0.8 (0. Vleeshouwers 1997.1) 8.8 (3. hairy bittercress.3) 8.Echinochloa crus-galli.4 (0.1) Johnsongrass 6.2 (2.0) Blackgrass 5. 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.2 (0.8) 11.4) Common purslane 5.3) Buckhorn plantain 7.5) 8. level of emergence we found was slightly greaterthan values scribed in the cited studies.9 (3. the effects of soil compaction were not emergence inhibition increases proportionately with depth examined. and above all.wild mustard.2) Large crabgrass 6. Datura stramonium.3 (0.2) 8.4 (0.3) 4.7 (0.7) Canada thistle 5.8 (0.3 (1.6) 12.soil compaction can act directly physical characteristics. .7) 6.6 (1.4) 7. Black.Digitaria sanguinalis. which were generally able to germinate and reach the soil Webb et al.9 (0.7) 7.cm)b Weedspeciesa 0 2 4 6 8 10 12 d Prostrate knotweed 6. and field bindweed.2 (1. common chickweed.3) 16.7 (2.2 (0.0) 13.8) 14.5 (0.3 (0.6) 10.

seed specific weight could play a major Common lambsquarters 0. Daturastramonium. g).052 role in allowing germination at a considerable distance be- Redroot pigweed 0.234 Barnyardgrass 0. Means are followed by their to be linked to increasing difficulty in gas exchange with relative standard errors. SD 0 UD 50 . 9070 T E T T IT T T T T0 90 : 0 _~~~~~ U) Tz3 + ~~ si g8 . the germination behavior of seeds buried at in- Velvetleaf 8.000-seedweight mancy seems to be mediated by lack of 02 (Benvenuti and Macchia 1995. A possible correlation between seed specific weight (Table knotweed.083 limited autotrophic phase (beginning of photosynthesis). This speculation was confirmed by noting that res- ure 2). Seeds germi- Prostrateknotweed 1.5 cutleafgeranium. 20J |80 T T 11?T?* ? 8 v 0 PL IU. Q 70 'C cn ?70~u) E - 60 Q ) 0 60 ~C.312 ygen deficiency (Al-Ani et al.commonpurslane. induction of seed dor- Weed species 1.velvetleaf.034 However. 1989). 100 100 T T a.052 In this perspective. . Table 2 shows the specific weight (re- Common chickweed 0.175 inversely correlated with depth (Drew 1990). given that oxygen concentration is Canada thistle 1. 1998) or by an increase in CO2 (Holm g 1972) deriving from seed metabolism.034 Noticeable variation was found. In particular. weed and common lambsquarters (both slightly below 0.008 Hairy bittercress 0. Jimsonweed 8.763 creasing depths may also be linked to seed energy reserves.192 + 0. 1985). TABLE 2.094 ing the soil surface that represents the target of the energy- Buckhorn plantain 0.803 ? 0. ranging from over 11 g in Common purslane 0.jimsonweed. intermediate values.750 + 0. and although it has not yet been fully clarified. prostrate Digitariasanguinalis.133 ? 0.075 nating at excessive depth would have little chance of reach- Black nightshade 0. ed by inducing depth-mediated dormancy.Spergulaarvensis.~~~ U) ~~~~~~20 10 0 0 POLLA PLA CI.143 Field bindweed 9. noncompacted soil conditions.Amaranthusretro.2) and depth-mediated 50% emergence inhibition was ex- flexus-.041 ferring to 1.0Q- ~~~~~ 30 ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ -3Q0 cn 20 . Vertical bars indicate standard errors of the mean.115 ? 0.Cardamine hirsuta.574 cies to allow activation of seed metabolism even under ox- Cutleaf geranium 2. largecrabgrass. Abutilontheophrasti.000-fold lower seed weight in hairy bittercress (both less than 0. Examination of seeds recov. The means across species are indicated by the solid line. all (approximately 85%) remained completely dormant (Fig.051 low the soil surface. specific weights in the upper range were blackgrass.Plantagolanceolata.41 + 0.Polygonumlaphathifolium. cornspurry. blacknightshade. Catchweed bedstraw 11.508 ? 0.858 + 0.911 looked that virtually the entire seed bank is subjected to Wild mustard 1.777 ? 1.088 ? 0.000 seeds) of each of the 20 species studied.424 + 0. to achieve germination percentages similar to values ob- served in unburied seeds.AR ALOMY ECHCG CHEAL 'ABUh GAAPPOROL SEM AWAE DATST DIOSA SORHA SINA SOLNI CARHI GERDI CONA SPRA Weed species FIGURE 2.fieldbindweed.885 + 0.Stellariamedia.645 ? 0. It should not be over- Blackgrass 1.203 some degree of hypoxia. 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. Chenopodiumal. Convolvolus arvensis.603 ? 0. Depth-mediated dormancy was reported by Wesson and Wareing (1969) as early as 1969.commonlambsquarters. Large crabgrass 0. Canadathistle.092 + 0.Galiumaparine. increasing soil depth.2 g) and in the lower range for redroot pig- bum.Solanumnigrum.: Buried weed seedling emergence * 533 .Alopecurusmyosuroides.847 because reserves are known to be crucial in some crop spe- Johnsongrass 5. john- songrass. 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. Cirsiumarvense-. recorded for field bindweed (9.373 ? 0.697 + 0. velvetleaf (8. Brassicakaber. Echinochloacrus-galli. hairybittercress.catchweedbedstraw.000-seed weight) tested in the trials. it appears 1.813 + 0.com. 0 cn E -o 0 - 0 U. Geranium dissectum. 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. jimsonweed (8. Portulacaoleracea. Corn spurry 0.buckhorn plantain.490 + 0.1 g).6 g). redrootpigweed.wildmustard. Weed species (listed as a function of decreasingsize. A statistically significant correlation (P < Benvenuti et al. and mon chickweed. Sorghumhalepense-.231 ? 0.7 g). Among a Barnyardgrass. so that germination was very limited (only 15%).007 catchweed bedstraw to a more than 1. plored (Figure 1).

sons. P. and L. A model of the effects of cultivation on the vertical distribution of weed seeds within the soil. S. M. Weed Sci. It is Western Nebraska. R.C. G. 1995. Eindhoven. Wilson. the pronounced seed longevity (Burnside et al. C. vel. E.) and broadleaf Weed Sci. 45:85-90. Weed biology: importance to weed management. Hypoxia effect on buried weed seed germination. achievingprecisionin seed dynamicsimulationwill it be- come possible to make weed control optimally effective 7My 0 while minimizingunjustifiedherbicideapplication. Agric. cation of herbicides. chem.. D. 0. 79:885-890. M. Mediterr. they clarify the complex ecophysiology of soil Cidecydan. This indicates that seed weight can represent a seed germination and emergence rates. Weed Res. been hypothesized further that the pronounced fluctuation Bhowmik. 123:252-256. Bruzau. R. Weed Sci. Taylor and Ten Broeck 1980. and K. Licor.e. and A. Fisher Scientific. a relation between seed longevity and unit phrasti Medicus. the like. Scopel. The annual dormancy cycle in buried weed seeds: a continuum. R. seeds. Leblank. 4 CoHort Software. Stefani. Sain-Ges.4-. G. Weed Res. 1996) could favor depth-mediated depth effects on redstem filaree (Erodiumcicutarum)emergence. 1992. weight has been suggested (Thompson et al.639 x+ 4. Radosevich. M. 1994. July-August 2001 . R. at least in the species tested. R. 1998). C. 1999). G. R. Phytochrome mediated germination control of Datura stramoniumL. Baskin. Ballare. M. 1997. J. Bond. An index for predicting (Figure 3).12 1 Filter paper. A. and S. or harvesting. 1997. worth pointing out that a seed unit weight effect has been Cardina. Seed bank production and seedbank dynamics in subthreshold velvetleaf (Abutilon theophrasti)populations. C.. dock (Rumex obtusijfoliusL. Germination. V. The present labo. R. Weed mination inhibition appears to be an important survival Sci.000 seed weight (g) FIGuRE 3.(jonsonngrass. in seed unit weight sometimes present even within the same Blackshaw. Datura stramoniumL. Macchia. Weed Sci. and S. M. First. J. 1997. beggarticks (Bidens tripartita) as affected by light and oxygen. Weed seedbank response to tillage. 1995. L. M.. 40:204-207.. Macchia. A. S. Macchia. This would Sci. Chancellor. MN 55419. and M. 7:560-569. J. production and some morphological characteristicsof Abutilon theo- In addition. 35:343-351. and C. 45:696-700. Benvenuti. Macchia. F. 56:777-788. 1996.). Effects of seed size on the depth inhibition of weed seed germination. and A. S. described in curly dock (Rumex crispusL. Ecology 70:227-232. dormancy as causes of failed emergence. Norquay.J. Philips TLF 20W/33. and SorghumhalepenseL. Sparrow.PA o o R 0. Soil temperature. Benvenuti. 0. Weed Technol.. Weed Sci. for identifying the Ball. J. Al-Ani. Cussans. n 0 1 2 3 4 5 6 7 8 9 10 11 12 3 Spectroradiometermodel 1800. 1996) that Benvenuti. 0. Plant Physiol. S. It could underlie Benvenuti. 1993). that involve or simulate seed bank dynamics.05) according to a second-degree equation was observed Alm. Minneapolis.J. Wax. Weisberg. valid tool. M. 45:349-356. and M.000-seed weight. 1993. Malloch 1982). ported for various crop species (Harper and Obeid 1967. emergence: an allometric relationship and some ecological implica- tions. and A. M. S. W. D. 1964. Seed longevity of 41 weed species buried 17 years in eastern and vorable ecological situations (Hodkinson et al. Effects of shade on re- leads to a persistent seed bank (Thompson 1987). S. It has Weed Res. Heather and Sieczka 1991. Photo- A relation between seed size and emergence has been re. J. Soil light penetration and dormancy of Jimsonweed Weaver 1980) and for wild species (Bond et al. 1.036 x2he0. Inc. 1990. Photobiol. and crop rotation sequence. W. L. This mechanism of ger. soil moisture and seed burial species (Milberg et al. 1992. The . Rersinbtenbra0et eesr ordc mrec 2 Fluorescent lamp. A. and H. 1998. Moss. J. P. NE. S.. mex obtusifolius. Cousens. But (Datura stramonium)seeds. Cumbenworth. C. Calculation of threshold temperature possible to distinguish between suicide germination and seed for the development of various weeds. and K.This will favor harmonizationof ecologicalconcernswith the eco- a 30 3456 C 08 nomic advantagesexpectedfrom modernagriculture. and M. Photomorphogenic processes in the agriculturalenvironment. Raymind. weed seed size may be correlated with resistance to unfa.05) and the corresponding R2 value are shown.75 15219. Only by C) 4-1 . Raudonius. Lincoln. Honig. FIUE 3. Sanchez. suggesting that Burnside. Germination ecophysiology of bur due to depth-imposed dormancy. 40:654-659. Malloch. A. 34:283-288. strategy. 1982.0 Netherlands. lihood of emergence and consequent crop invasion).. growth and competitive ability of Rumexcrispusand Ru- demons. 38:199-205. Maze 1999. Regression between burial depth necessary to reduce emergence of the 20 weed species tested to 50% and corresponding 1. ratory tests demonstrated that failed emergence was mainly Benvenuti. Bioscience 35:492-498. Macchia. withi few excepntions. Emergence of weed seedlings in the field and the effects of different frequencies of cultivation. Weed Sci. 44:74-86. 1. 1985. 1996. Pradett. 43:389-393. it was not Benvenuti. 45:61-66. Baskin. Literature Cited The equation (significant for P > 0. Weed dormancy heteroblasty of buried weed seeds. and S. Temporal changes in velveatleaf influence emerged seedling competitiveness (Cidecydan and (Abutilon theophrasti)seed dormancy. Second. where seed size was shown to Cardina. Seed size and seedling tion ability despite agronomic disturbances such as appli. Brain. herbicides. allowing seed bank perpetuation. tillage. Oecologia 120:132-136.traqtethat. Pittsburgh. E. 30:61-70. Pers. A. H. 1993. Watchman No. and M. E. respiration and adenylate charge of seeds at various oxygen pressures. 1992... Ef- fects of depth of seed burial and soil aggregate of Alopecurusmyosu- 534 * Weed Science 49. Pages 599-606 in Pro- Results obtained in this study are of interest for two rea- ceedings of the Seventh British Weed Control Conference. Hubbard. 1997.. W Stoller. they germination. emergence profile of the active soil seed bank (i. and D. 1985. P. S. 0 0) Sources of Materials eq ( y=. because such trials were conducted in the field. Brighton. ensure that buried dormant seeds maintain their germina. Weed Res.

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