Promotion of seed germination in Echinochloa crus-galli with atmospheric air plasma

Promotion of seed germination in Echinochloa crus-galli with atmospheric air plasma | bioRxiv /* */ /* */ <!-- <!-- /*! * yepnope1.5.4 * (c) WTFPL, GPLv2 */ (function(a,b,c){function d(a){return"[object Function]"==o.call(a)}function e(a){return"string"==typeof a}function f(){}function g(a){return!a||"loaded"==a||"complete"==a||"uninitialized"==a}function h(){var a=p.shift();q=1,a?a.t?m(function(){("c"==a.t?B.injectCss:B.injectJs)(a.s,0,a.a,a.x,a.e,1)},0):(a(),h()):q=0}function i(a,c,d,e,f,i,j){function k(b){if(!o&&g(l.readyState)&&(u.r=o=1,!q&&h(),l.onload=l.onreadystatechange=null,b)){"img"!=a&&m(function(){t.removeChild(l)},50);for(var d in y[c])y[c].hasOwnProperty(d)&&y[c][d].onload()}}var j=j||B.errorTimeout,l=b.createElement(a),o=0,r=0,u={t:d,s:c,e:f,a:i,x:j};1===y[c]&&(r=1,y[c]=[]),"object"==a?l.data=c:(l.src=c,l.type=a),l.width=l.height="0",l.onerror=l.onload=l.onreadystatechange=function(){k.call(this,r)},p.splice(e,0,u),"img"!=a&&(r||2===y[c]?(t.insertBefore(l,s?null:n),m(k,j)):y[c].push(l))}function j(a,b,c,d,f){return q=0,b=b||"j",e(a)?i("c"==b?v:u,a,b,this.i++,c,d,f):(p.splice(this.i++,0,a),1==p.length&&h()),this}function k(){var a=B;return a.loader={load:j,i:0},a}var l=b.documentElement,m=a.setTimeout,n=b.getElementsByTagName("script")[0],o={}.toString,p=[],q=0,r="MozAppearance"in l.style,s=r&&!!b.createRange().compareNode,t=s?l:n.parentNode,l=a.opera&&"[object Opera]"==o.call(a.opera),l=!!b.attachEvent&&!l,u=r?"object":l?"script":"img",v=l?"script":u,w=Array.isArray||function(a){return"[object Array]"==o.call(a)},x=[],y={},z={timeout:function(a,b){return b.length&&(a.timeout=b[0]),a}},A,B;B=function(a){function b(a){var a=a.split("!"),b=x.length,c=a.pop(),d=a.length,c={url:c,origUrl:c,prefixes:a},e,f,g;for(f=0;f<d;f++)g=a[f].split("="),(e=z[g.shift()])&&(c=e(c,g));for(f=0;f<b;f++)c=x[f](c);return c}function g(a,e,f,g,h){var i=b(a),j=i.autoCallback;i.url.split(".").pop().split("?").shift(),i.bypass||(e&&(e=d(e)?e:e[a]||e[g]||e[a.split("/").pop().split("?")[0]]),i.instead?i.instead(a,e,f,g,h):(y[i.url]?i.noexec=!0:y[i.url]=1,f.load(i.url,i.forceCSS||!i.forceJS&&"css"==i.url.split(".").pop().split("?").shift()?"c":c,i.noexec,i.attrs,i.timeout),(d(e)||d(j))&&f.load(function(){k(),e&&e(i.origUrl,h,g),j&&j(i.origUrl,h,g),y[i.url]=2})))}function h(a,b){function c(a,c){if(a){if(e(a))c||(j=function(){var a=[].slice.call(arguments);k.apply(this,a),l()}),g(a,j,b,0,h);else if(Object(a)===a)for(n in m=function(){var b=0,c;for(c in a)a.hasOwnProperty(c)&&b++;return b}(),a)a.hasOwnProperty(n)&&(!c&&!--m&&(d(j)?j=function(){var a=[].slice.call(arguments);k.apply(this,a),l()}:j[n]=function(a){return function(){var b=[].slice.call(arguments);a&&a.apply(this,b),l()}}(k[n])),g(a[n],j,b,n,h))}else!c&&l()}var h=!!a.test,i=a.load||a.both,j=a.callback||f,k=j,l=a.complete||f,m,n;c(h?a.yep:a.nope,!!i),i&&c(i)}var i,j,l=this.yepnope.loader;if(e(a))g(a,0,l,0);else if(w(a))for(i=0;i (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0];var j=d.createElement(s);var dl=l!='dataLayer'?'&l='+l:'';j.src='//www.googletagmanager.com/gtm.js?id='+i+dl;j.type='text/javascript';j.async=true;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-M677548'); Skip to main content Home About Submit ALERTS / RSS Search for this keyword Advanced Search New Results Promotion of seed germination in Echinochloa crus-galli with atmospheric air plasma Yuto Nakamura , Yu Ohnishi , Kazuma Oshimo , Rei Hiramatsu , Masako Shindo , Tohru Tominaga , Satoshi Terabayashi , Daisuke Takagi , View ORCID Profile Mitsuhiro Matsuo doi: https://doi.org/10.1101/2025.11.14.688452 Yuto Nakamura 1 Department of Applied Biological Science, Faculty of Agriculture, Setsunan University , Hirakata, Osaka, Japan Find this author on Google Scholar Find this author on PubMed Search for this author on this site Yu Ohnishi 1 Department of Applied Biological Science, Faculty of Agriculture, Setsunan University , Hirakata, Osaka, Japan Find this author on Google Scholar Find this author on PubMed Search for this author on this site Kazuma Oshimo 1 Department of Applied Biological Science, Faculty of Agriculture, Setsunan University , Hirakata, Osaka, Japan Find this author on Google Scholar Find this author on PubMed Search for this author on this site Rei Hiramatsu 1 Department of Applied Biological Science, Faculty of Agriculture, Setsunan University , Hirakata, Osaka, Japan Find this author on Google Scholar Find this author on PubMed Search for this author on this site Masako Shindo 2 Department of Applied Biological Science, Osaka Institute of Technology , Osaka, Japan Find this author on Google Scholar Find this author on PubMed Search for this author on this site Tohru Tominaga 3 Kyoto University , Kyoto, Japan Find this author on Google Scholar Find this author on PubMed Search for this author on this site Satoshi Terabayashi 1 Department of Applied Biological Science, Faculty of Agriculture, Setsunan University , Hirakata, Osaka, Japan Find this author on Google Scholar Find this author on PubMed Search for this author on this site Daisuke Takagi 1 Department of Applied Biological Science, Faculty of Agriculture, Setsunan University , Hirakata, Osaka, Japan Find this author on Google Scholar Find this author on PubMed Search for this author on this site Mitsuhiro Matsuo 1 Department of Applied Biological Science, Faculty of Agriculture, Setsunan University , Hirakata, Osaka, Japan Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Mitsuhiro Matsuo For correspondence: mitsuhiro.matsuo{at}setsunan.ac.jp Abstract Full Text Info/History Metrics Preview PDF Abstract Plasma technology has attracted increasing attention for potential agricultural applications, and it has been investigated for decades particularly for its ability to promote seed germination. However, most studies have focused on crop species, and little is known about its potential use for weed management. In this study, we examined whether air plasma influences the germination of Echinochloa crus-galli , a major paddy field weed. When seeds were cultivated without stratification, direct irradiation with air plasma did not enhance germination. In contrast, once dormancy was partially released by cold stratification, plasma treatment significantly promoted germination. Interestingly, the effect disappeared after prolonged chilling, when dormancy was completely released. These results suggest that plasma promotes germination by acting on the dormancy-breaking process. In addition, plasma-activated water (PAW) also stimulated germination, although its effect was weaker than that of gaseous plasma. Overall, this study demonstrates that air plasma can promote the germination of grass weed seeds, providing a potential strategy for weed control by inducing germination prior to crop seeding. Introduction Weeds are considered major pests that significantly reduce crop yields ( Chauhan, 2020 ; Oerke, 2006 ). To maintain high levels of crop productivity, effective weed management strategies are essential. A key trait that leads to the persistence of weeds and increases the burden on farmers is their year-round emergence. This trait is primarily attributed to seed dormancy, seed longevity and responsiveness to environmental changes. Dormancy serves as an adaptive mechanism that prevents germination during unfavorable seasons or under suboptimal environmental conditions (Finch Savage and Leubner Metzger, 2006). During this period, seeds remain metabolically inactive yet viable. In addition, many weed seeds exhibit long-term viability, persisting in the soil seedbank for several years or even decades ( Conn et al., 2006 ). These seeds may germinate opportunistically when environmental conditions become favorable, complicating weed control and increasing labor and management costs. To mitigate the difficulty of weed management, the stale seedbed technique has been developed as a cultural weed management strategy ( Gallandt, 2006 ; Travlos et al., 2020 ). This method promotes weed seed germination before crop sowing through soil disturbance practices such as shallow tillage. It exploits the tendency of many weed seeds to germinate in response to light exposure, temperature fluctuations, and moisture availability. Once germinated, these seedlings can be eliminated by mechanical, thermal, or chemical means prior to planting the main crop, thereby reducing weed pressure during the critical early stages of crop development. Furthermore, in the case of Striga spp. (witchweed), which are parasitic weeds of major concern in sub-Saharan Africa, synthetic strigolactones, which are compounds that stimulate Striga germination, can be applied in the absence of a host plant ( Zwanenburg et al., 2016 ). As a result, Striga seeds germinate and subsequently die due to the lack of a suitable host. This “suicidal germination” approach is conceptually similar to the stale seedbed technique and highlights the broader potential of exploiting seed biology in weed management. These approaches not only reduce the frequency of weeding during crop cultivation but also help deplete the weed seedbank in the soil. Continued development and refinement of such techniques are essential for sustainable agricultural systems. In recent decades, the application of plasma technology to plant science and agriculture has been intensively studied ( Bourke et al., 2018 ). One important application is seed processing technology, in which plasma treatment is used to promote seed germination ( Grainge et al., 2022 ; Waskow et al., 2021 ). Plasma treatment has been suggested to induce alterations in the physical and biochemical properties of seeds, thereby influencing germination ( Waskow et al., 2021 ). In particular, air plasma abundantly generate reactive oxygen species (ROS) and reactive nitrogen species (RNS), which are considered as key agents because they have been reported as signaling molecules regulating seed germination ( Bykova and Igamberdiev, 2024 ; Puglia, 2024 ). The positive effects of air plasma on seed germination have been reported across a wide range of species, including not only angiosperms but also gymnosperms ( Priatama et al., 2022 ; Waskow et al., 2021 ). However, most studies have focused on crop species, whereas knowledge related to weed management remains limited. Bukhori et al. (2024) reported that high-voltage plasma treatment reduced the germination rate of mustard seeds and suggested that plasma technology could potentially be applied to control weed seed germination. However, their study did not directly involve weed species and did not examine the potential application of the promotive effects of plasma on seed germination in the context of weed control. Theoretically, promoting germination with plasma treatment, followed by the elimination of seedlings before crop planting, could be an effective strategy for weed control and soil seedbank management. In this study, we investigated the effect of plasma on the germination of Echinochloa crus-galli , one of the most problematic weeds in both paddy and upland fields ( Tian et al., 2020 ; Zhang et al., 2021 ), and showed that plasma could induce the germination in a manner dependent on dormancy status. Our findings suggest that plasma could serve as a novel tool to stimulate weed seed germination for pre-sowing weeding and soil seedbank regulation. Material and methods Plant material and cultivation Seeds of Echinochloa crus-galli were collected from a paddy field in Higae-cho, Keihoku, Ukyo-ku, Kyoto City, Japan (35.1819°N, 135.6665°E). The collected seeds were propagated in a controlled-environment growth chamber to increase the seed stock. Seeds obtained from the cultivations were used for the germination experiments. Plants were grown under controlled conditions of 25 °C with a 14-hour light and 10-hour dark photoperiod. Plasma irradiation and construction of plasma-activated water Atmospheric pressure air plasma was generated using a dielectric barrier discharge (DBD) system with pen-type plasma device. The system was constructed of power supply (LHV-13AC, LOGY ELECTRIC CO.,LTD., Tokyo, Japan), an air-pump (e-air 2000SB, Gex, Osaka, Japan) and a handmade pen-type electrode. The electrode was constructed by placing a stainless wire mesh inside a ceramic tube (4 mm inner diameter, 6 mm outer diameter) as the high-voltage electrode, and attaching a copper sheet on the outside of the ceramic tube as the grounded electrode. The air plasma was applied to the seeds of Echinochloa crus-galli for 15 minutes. Plasma-activated water (PAW) was prepared by introducing the plasma into 1 L of deionized water for 3 hours at 22C°C, with a flow rate of 0.5 L/min. Deionized water aerated without plasma for 3 hours were used as control. PAW and control water was applied to the double ring filter paper (99-191-090, Cytiva, USA), where the seeds were sown. Germination test 30 seeds were sown on filter paper. Stratification of the seeds were performed at 4 C°C under darkness for the specified periods. And then the they were transferred to light condition at 22C°C. Germinated seeds were counted and the average germination ratio and standard deviation were calculated from three independent experiments. Measurements of nitrate, nitrite and hydrogen peroxide NOCC and NO 2 C were measured using the spectroscopic method described by according to ( Hachiya and Okamoto, 2017 ). H 2 O 2 was measured using Amplite®Fluorimetric Hydrogen Peroxide Assay Kit (Red fluorescence, AAT Bioquest, Pleasanton, CA. USA) according to the manufacture’s instruction. Results Pen-type DBD plasma irradiation system and the Plasma-activated water In this study, air plasma was generated using a handmade pen-type dielectric barrier discharge (DBD) plasma system and used for seed treatment experiments. The system delivered air (0.7 L minC¹) into the nozzle electrode via an air pump, where it was converted into air plasma by applying a high voltage (2.3 W, Vpp 9 kV, 13.7 kHz) generated from an AC power supply ( Figure 1A, B ). The generated plasma, which emitted purple light ( Figure 1C ), was either directly irradiated onto Echinochloa crus-galli seeds or bubbled into water to prepare plasma-activated water (PAW) ( Figure 1B ). Download figure Open in new tab Figure 1. Generation of atmospheric air plasma and plasma-activated water (PAW). A) Overview of the handmade atmospheric air plasma generation device. The device mainly consists of four parts: a power supply, an air pump, and a nozzle electrode. B) Schematic illustration of the air plasma treatment. Air delivered by the pump to the nozzle electrode is ionized under high voltage to generate plasma. The ejected plasma was either directly irradiated onto Echinochloa crus-galli seeds or bubbled into deionized water to produce plasma-activated water (PAW). C) Air plasma emitting purple light inside the nozzle electrode. Plasma irradiation promotes the germination of Echinochloa crus-galli dependent on dormancy break Echinochloa crus-galli seeds have deep dormancy, which can be broken by cold treatment (Honek et al., 1999). To explore the effect of air plasma on the germination of Echinochloa crus-galli seeds, seeds were treated with air plasma and incubated either with or without cold treatment ( Figure 2A–D ). The air plasma treatment significantly promoted germination when seeds were stratified with one month of cold treatment ( Figure 2C ). However, no promotive effect of air plasma was observed in seeds incubated without cold treatment or with only a short, one-week cold treatment, which was not sufficient to break dormancy and resulted in a low germination rate ( Figure 2A , B). Furthermore, most seeds were germinated after two months of cold stratification and there was less difference between the plasma treated seeds and the control seeds ( Figure 2D ). These results suggest that air-plasma should promote the germination of Echinochloa crus-galli but the positive effect should occur during the periods when the dormancy is released. Download figure Open in new tab Figure 2. Effect of plasma irradiation on the germination of Echinochloa crus-galli Echinochloa crus-galli seeds were directly irradiated with atmospheric air plasma, and then sown on wet filter paper. Plasma-irradiated and control seeds were transferred from stratification conditions (4 °C, darkness) to germination conditions (22 °C, continuous light), and then cultivated for 2–4 months. The horizontal axis indicates the cultivation period under germination conditions, and the vertical axis represents the percentage of germinated seeds. A) No stratification. B–D) Seeds stratified for 1 week (B), 1 month (C), and 2 months (D), respectively. Data are presented as means ± standard deviations. Asterisks indicate significant differences according to the t -test ( p < 0.05). Plasma-activated water promote the germination of Echinochloa crus-galli When considering the application of the germination-promoting effect of plasma in agriculture, it would be practical to use air plasma in the form of plasma-activated water (PAW), produced by infusing air plasma into water. We examined whether PAW has a similar germination-promoting effect ( Figure 3 ). When the seeds were placed on filter paper soaked in PAW, no promotion of the germination was observed in dormant seeds without cold treatment ( Figure 3A ). After 2 weeks cold treatment, in a part of the PAW treated seeds, germination was promoted ( Figure 3B ), although most of the seeds remained to be ungerminated. In contrast to direct plasma irradiation of Figure 2C , there was less promotion in 1 month cold treated seeds ( Figure 3C ). For 2 months cold treated seed, whose dormancy should have been broken, no promotion effect by PAW was detected as well as Figure 2D . These results suggest that the liquid form of air plasma should have germination promotion effect of Echinochloa crus-galli seeds during seed dormancy breaking periods. However the effect should be less compared to the direct plasma irradiation. Download figure Open in new tab Figure 3. Effect of plasma-activated water (PAW) on the germination of Echinochloa crus-galli Echinochloa crus-galli seeds were treated with PAW or air-bubbled water (control) and then sown on wet filter paper. PAW-treated and control seeds were transferred from stratification conditions (4 °C, darkness) to germination conditions (22 °C, continuous light), and then cultivated for more than one month. The horizontal axis indicates the cultivation period under germination conditions, and the vertical axis represents the percentage of germinated seeds. A) No stratification. B–D) Seeds stratified for 2 weeks (B), 1 month (C), and 2 months (D), respectively. Data are presented as means ± standard deviations. An asterisk indicates a significant difference according to the t -test ( p < 0.05). Plasma-activated water contain nitrate and hydrogen peroxide It was reported that PAW of air could contain nitrate (NO 3 - ), nitrite (NO 2 - ) and hydrogen peroxide (H 2 O 2 ) ( Sivachandiran and Khacef, 2017 ). These molecules have been shown to have the effect of germination promotion ( Bethke et al., 2006 ; Duermeyer et al., 2018 ; Wojtyla et al., 2016 ). To unveil how much these molecules are included in the PAW generated by pen-type plasma device in this study, we measured the amount of NO 3 - , NO 2 - and H 2 O 2 in the PAW. The analysis showed that the PAW contained 1.9 ×10 2 μM NO 3 - and 9.4 ×10 -1 μM H 2 O 2 , but no NO - was detected ( Table 1 ). View this table: View inline View popup Download powerpoint Table 1. Concentrations of NO 3 - , NO 2 - and H 2 O 2 in Plasma-activated water (PAW) PAW was prepared as described in Material and Methods section, and air-bubbled deionized water was used as control. nd, below detection limit. Data are presented as means ± standard deviations. n ≥ 3. Asterisk indicates significant differences compared with control according to t -test (p < 0.05). Discussion It has been reported that atmospheric air plasma has positive effect on plant germination and the characteristics is expected to apply to agriculture. However there is few knowledge for utilization of the positive effect into weeding. In this study, we examined whether air plasma has germination promotion effect on Echinochloa crus-galli , one of the major weed in paddy field world wide, and showed that the plasma and plasma-activated water has the germination promotion effect in this weed. Similar to many other weed species, Echinochloa crus-galli seeds exhibit long-term dormancy, and their germination largely depends on the degree of dormancy. To examine the effect of air plasma on germination and dormancy, we treated the seeds with the plasma and evaluated germination after different periods of cold stratification. Our results showed that the promotive effect on germination was evident only during the dormancy-breaking period in Echinochloa crus-galli ( Figure 2C ). In addition, no promotion was observed in seeds without cold stratification ( Figure. 2A ), and the effect disappeared once dormancy was fully released ( Figure 2D ). These results indicate that the plasma alone does not fully break dormancy but rather accelerates the dormancy-breaking process in Echinochloa crus-galli . Seed dormancy maintenance and release are regulated by highly complex molecular systems ( Bykova and Igamberdiev, 2024 ; Puglia, 2024 ; Sajeev et al., 2024 ). Air plasma may act on regulatory factors to attenuate seed dormancy in Echinochloa crus-galli . In this study, we showed that plasma-activated water (PAW) also promoted seed germination in Echinochloa crus-galli , but its effect was markedly weaker than that of direct plasma irradiation on seeds. This may be because the amount of germination-activating molecules in PAW is insufficient. Atmospheric air plasma is known to contain various reactive nitrogen and oxygen species (RNS and ROS) in abundance ( Cimerman and Hensel, 2023 ). Among the RNS, nitogen oxide (NO) has been reported to promote seed germination as signal molecule ( Arc et al., 2013 ). NO appears to be major factor responsible for the germination-promoting effect of air plasma. However NO is unstable and is rapidly oxdized into NO 2 - and NO - in water. NO 3 - also promotes germination and is likely one of key molecules regulating germination in PAW. Huang et al (2025) reported that the molecular mechanism by which NO 3 - induces germination is distinct from that of NO ( Huang et al., 2025 ). In addition, Bethke et al. (2006) demonstrated that a much higher concentration of NO - is required to break seed dormancy compared with NO in Arabidopsis thaliana . They showed that 100 ppm (≈4 μM) NO could induce about 40% seed germination, whereas approximately 1.0 mM NO 3 - was needed to achieve similar germination rate ( Bethke et al. 2006 ). In this study, 0.2 mM NO 3 - was detected in PAW. This concentration is far lower than the NO 3 - concentration, which induce germination, previously reported in Arabidopsis thaliana ( Bethke et al. 2006 ) and Echinochloa colona ( Picapietra and Acciaresi, 2022 ). The weak germination-promoting effect of PAW may be attributed to its low NO 3 - concentration. In addition to NO - , H O, which is one of the stable ROS in water, was also detected in PAW ( Table 1 ), and it has been reported to promote seed germination ( Ogawa and Iwabuchi, 2001 ). However, the concentration (0.9 μM, Table 1 ) is about two order of maginitude lower than that reported in other studies ( Barba-Espín et al., 2012 ; Ogawa and Iwabuchi, 2001 ). Therefore the limitted effect of PAW may be due to the low concentration of these molecules. Increasing the concentration of NO 3 - and H 2 O 2 concentration by optimizing the PAW generation system may enhance its germination-promoting effect, although this remains to be investigated in future work. The stale seedbed technique, which promotes weed germination and subsequent weeding, is widely used in agricultural practice, and its improvement could contribute to future sustainable agriculture. Our results showed that the germination of a grass weed could be promoted by plasma and PAW, suggesting that plasma technology has the potential to enhance and extend chemical-based stale seedbed techniques for weed management. Notably, the germination-promoting effect of plasma has been reported across a wide range of plant species ( Priatama et al., 2022 ; Waskow et al., 2021 ). Its application to weeding does not seem to be limited to Echinochloa crus-galli , but could potentially cover diverse species. Furthermore, air plasma and PAW can be generated from air, water and household electricity using a simple device composed of a power supply, air pump and pen-type electrode. The simplicity of this device could enable on-site production of plasma and PAW, rather than relying on an industrial setting. In addition, plasma generates multiple molecular species, including NO, NOCC, and HCOC, which promote germination in a single step. The composition of air plasma can be adjusted depending on the plasma device and the generation conditions ( Cimerman and Hensel, 2023 ; Klenivskyi et al., 2024 ). This flexibility allows the plasma composition to be tailored according to seed species, physiological state, and environmental conditions for agricultural applications. These characteristics may offer new alternatives in agricultural practice. However, the effects of plasma on germination and plant growth also depend on seed age and physiological state ( Attri et al., 2021 ). Under natural field conditions, seeds in the soil exhibit diverse physiological states. Therefore, numerous factors need to be considered when applying plasma and PAW treatments to weed seed germination. Further investigations integrating plasma technology with plant science and agronomy are required to assess the practical feasibility and potential applications of plasma technology for weed control. Conflicts of Interest The authors declare no conflicts of interest. Acknowledgement This study was supported by JSPS KAKENHI Grant Numbers JP23K03371. We gratefully thank Prof. Junichi Obokata and Prof. Junshi Yazaki for their critical and productive discussions. Funder Information Declared Japan Society for the Promotion of Science (JSPS) , JP23K03371 Footnotes Zhongcunyouren02{at}gmail.com , butterfly0024yu{at}outlook.jp , 22b012ok{at}edu.setsunan.ac.jp , 22b072hr{at}edu.setsunan.ac.jp , masako.shindo{at}oit.ac.jp , tominaga.tohru.5e{at}kyoto-u.jp , satoshi.terabayashi{at}setsunan.ac.jp , daisuke.takagi{at}setsunan.ac.jp , mitsuhiro.matsuo{at}setsunan.ac.jp References ↵ Arc , E. , Galland , M. , Godin , B. , Cueff , G. , Rajjou , L ., 2013 . Nitric oxide implication in the control of seed dormancy and germination . Front. Plant Sci . 4 . doi: 10.3389/fpls.2013.00346 OpenUrl CrossRef ↵ Attri , P. , Ishikawa , K. , Okumura , T. , Koga , K. , Shiratani , M. , Mildaziene , V ., 2021 . 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Share Promotion of seed germination in Echinochloa crus-galli with atmospheric air plasma Yuto Nakamura , Yu Ohnishi , Kazuma Oshimo , Rei Hiramatsu , Masako Shindo , Tohru Tominaga , Satoshi Terabayashi , Daisuke Takagi , Mitsuhiro Matsuo bioRxiv 2025.11.14.688452; doi: https://doi.org/10.1101/2025.11.14.688452 Share This Article: Copy Citation Tools Promotion of seed germination in Echinochloa crus-galli with atmospheric air plasma Yuto Nakamura , Yu Ohnishi , Kazuma Oshimo , Rei Hiramatsu , Masako Shindo , Tohru Tominaga , Satoshi Terabayashi , Daisuke Takagi , Mitsuhiro Matsuo bioRxiv 2025.11.14.688452; doi: https://doi.org/10.1101/2025.11.14.688452 Citation Manager Formats BibTeX Bookends EasyBib EndNote (tagged) EndNote 8 (xml) Medlars Mendeley Papers RefWorks Tagged Ref Manager RIS Zotero Tweet Widget Facebook Like Google Plus One Subject Area Plant Biology Subject Areas All Articles Animal Behavior and Cognition (7635) Biochemistry (17690) Bioengineering (13892) Bioinformatics (41936) Biophysics (21451) Cancer Biology (18588) Cell Biology (25499) Clinical Trials (138) Developmental Biology (13378) Ecology (19899) Epidemiology (2067) Evolutionary Biology (24320) Genetics (15609) Genomics (22506) Immunology (17736) Microbiology (40394) Molecular Biology (17181) Neuroscience (88603) Paleontology (666) Pathology (2832) Pharmacology and Toxicology (4824) Physiology (7641) Plant Biology (15152) Scientific Communication and Education (2045) Synthetic Biology (4294) Systems Biology (9825) Zoology (2271)