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",isbn:"978-1-83962-718-7",printIsbn:"978-1-83962-717-0",pdfIsbn:"978-1-83962-754-5",doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!0,hash:"4df95c7f295de7f6003e635d9a309fe9",bookSignature:"Dr. Yajuan Zhu, Dr. Qinghong Luo and Dr. Yuguo Liu",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/8969.jpg",keywords:"Water Cycle, Water Use Strategy, Vegetation Dynamics, Plant Community, Precipitation, Carbon Emission, Soil Respiration, Autotrophic Respiration, Algae Crust, Wind, Temperature, Vegetation Stability",numberOfDownloads:null,numberOfWosCitations:0,numberOfCrossrefCitations:null,numberOfDimensionsCitations:null,numberOfTotalCitations:null,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"January 26th 2021",dateEndSecondStepPublish:"February 23rd 2021",dateEndThirdStepPublish:"April 24th 2021",dateEndFourthStepPublish:"July 13th 2021",dateEndFifthStepPublish:"September 11th 2021",remainingDaysToSecondStep:"a month",secondStepPassed:!1,currentStepOfPublishingProcess:2,editedByType:null,kuFlag:!1,biosketch:"Dr. Zhu holds a Ph.D. in Ecology and is currently an Associate Research Professor at the Chinese Academy of Forestry at the Institute of Desertification Studies, she has led a number of national projects while working there.",coeditorOneBiosketch:"Dr. Luo holds a Ph.D. in Physical Geography and is currently a Research Professor at the Institute of Afforestation and Sand Control, Xinjiang Academy of Forestry. She is a holder of several technological patents in her area of research.",coeditorTwoBiosketch:"Dr. Liu holds a Ph.D. in Ecology and is currently an Assistant Professor at the Institute of Desertification Studies, Chinese Academy of Forestry. He has published several international works that have been recognized.",coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"180427",title:"Dr.",name:"Yajuan",middleName:null,surname:"Zhu",slug:"yajuan-zhu",fullName:"Yajuan Zhu",profilePictureURL:"https://mts.intechopen.com/storage/users/180427/images/system/180427.jpg",biography:"Dr. Yajuan Zhu obtained her Bachelor's degree in Agriculture from Northwest Agriculture and Forestry University in 2002 and PhD in Ecology from Chinese Academy of Sciences in 2007. She was a postdoctoral fellow working on the topic of land desertification control in the Research Institute of Forestry, Chinese Academy of Forestry, followed by her appointment as an Assistant Professor at the Institute of Desertification Studies, Chinese Academy of Forestry and currently she is an Associate Research Professor at the same institute. She is a Master's supervisor with interests in plant ecology in deserts, biodiversity, stable isotope ecology, isohydrology and desertification control.",institutionString:"Chinese Academy of Forestry",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"1",totalChapterViews:"0",totalEditedBooks:"0",institution:{name:"Chinese Academy of Forestry",institutionURL:null,country:{name:"China"}}}],coeditorOne:{id:"340564",title:"Dr.",name:"Qinghong",middleName:null,surname:"Luo",slug:"qinghong-luo",fullName:"Qinghong Luo",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0033Y000032N5e7QAC/Profile_Picture_1605773886590",biography:"Dr. Qinghong Luo holds a Master's degree from Life Science College, Shihezi University (2006) and PhD in Physical geography from Xinjiang Ecology and Geography Institute, Chinese Academy of Sciences (2018). She was initially an Assistant Research Professor at Institute of Afforestation and Sand Control, Xinjiang Academy of Forestry, after an Associate Research Professor and currently she is a Research Professor at the same institute. Her research interests include desert vegetation dynamics, plant-soil interaction and desertification control among others. She has participated in a number of funded and non funded projects and is a holder of several patents.",institutionString:"Chinese Academy of Forestry",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"0",totalChapterViews:"0",totalEditedBooks:"0",institution:{name:"Chinese Academy of Forestry",institutionURL:null,country:{name:"China"}}},coeditorTwo:{id:"340567",title:"Dr.",name:"Yuguo",middleName:null,surname:"Liu",slug:"yuguo-liu",fullName:"Yuguo Liu",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0033Y000032N5hEQAS/Profile_Picture_1605774524148",biography:"Dr. Yuguo Liu obtained his bachelor's degree, majoring in Environmental Sciences from Inner Mongolia University in 2007 and doctoral degree, majoring in Ecology from Institute of Botany, the Chinese Academy of Sciences in 2013. He has been working as an Assistant Professor at the Institute of Desertification Studies, Chinese Academy of Forestry ever since. His research interests include ecological protection and restoration of fragile areas, and karst vegetation and rocky desertification control.",institutionString:"Chinese Academy of Forestry",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"0",totalChapterViews:"0",totalEditedBooks:"0",institution:{name:"Chinese Academy of Forestry",institutionURL:null,country:{name:"China"}}},coeditorThree:null,coeditorFour:null,coeditorFive:null,topics:[{id:"10",title:"Earth and Planetary Sciences",slug:"earth-and-planetary-sciences"}],chapters:null,productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"},personalPublishingAssistant:{id:"194667",firstName:"Marijana",lastName:"Francetic",middleName:null,title:"Ms.",imageUrl:"https://mts.intechopen.com/storage/users/194667/images/4752_n.jpg",email:"marijana@intechopen.com",biography:"As an Author Service Manager my responsibilities include monitoring and facilitating all publishing activities for authors and editors. 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Venkateswarlu",coverURL:"https://cdn.intechopen.com/books/images_new/371.jpg",editedByType:"Edited by",editors:[{id:"58592",title:"Dr.",name:"Arun",surname:"Shanker",slug:"arun-shanker",fullName:"Arun Shanker"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"878",title:"Phytochemicals",subtitle:"A Global Perspective of Their Role in Nutrition and Health",isOpenForSubmission:!1,hash:"ec77671f63975ef2d16192897deb6835",slug:"phytochemicals-a-global-perspective-of-their-role-in-nutrition-and-health",bookSignature:"Venketeshwer Rao",coverURL:"https://cdn.intechopen.com/books/images_new/878.jpg",editedByType:"Edited by",editors:[{id:"82663",title:"Dr.",name:"Venketeshwer",surname:"Rao",slug:"venketeshwer-rao",fullName:"Venketeshwer Rao"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"4816",title:"Face Recognition",subtitle:null,isOpenForSubmission:!1,hash:"146063b5359146b7718ea86bad47c8eb",slug:"face_recognition",bookSignature:"Kresimir Delac and Mislav Grgic",coverURL:"https://cdn.intechopen.com/books/images_new/4816.jpg",editedByType:"Edited by",editors:[{id:"528",title:"Dr.",name:"Kresimir",surname:"Delac",slug:"kresimir-delac",fullName:"Kresimir Delac"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"49512",title:"Frankia as a Biodegrading Agent",doi:"10.5772/61825",slug:"frankia-as-a-biodegrading-agent",body:'Frankia are filamentous nitrogen-fixing Gram-positive actinobacteria that are found as free-living microbes in the soil and in symbiotic associations with actinorhizal plants [1-5]. These bacteria fix nitrogen by converting atmospheric N2 into biologically useful ammonia and supply the host plants with a source of reduced nitrogen. Frankia are developmentally complex and form three cell types: vegetative hyphae, spores located in sporangia, and vesicles. Hyphae are septate structures and form the growing state of this microbe. Under appropriate conditions, either terminal or intercalary multilocular sporangia are produced and contain many spores. When mature, the spores are released from the sporangia. The spores are presumed to aid in the survival and dispersal of Frankia in the environment. Vesicles are produced under nitrogen-limited conditions and consist of unique lipid-enveloped cellular structures that contain the enzymes responsible for nitrogen fixation. Thus, vesicles act as specialized structures for the nitrogen fixation process. Frankia are able to establish symbiotic nitrogen-fixing associations with over 220 species of woody dicotyledonous plants, termed actinorhizal plants, that are found in eight families of angiosperms [1, 3-6]. The symbiosis with Frankia allows these actinorhizal host plants to colonize nutrient-poor soil and harsh environments. Actinorhizal plants have been used to recolonize and reclaim industrial wastelands and environments contaminated with heavy metals and toxic aromatic hydrocarbon [7-15]. The metabolic potential of these bacteria has only recently been investigated in the context of bioremediation [16-18].
Based on phylogenetic analysis, Frankia strains have been classified into four main lineages [19-23]. Members of lineage 1 are found infective on host plants of the Betulaceae (Alnus), Myricaceae, and Casuariaraceae families, while lineage 2 represents strains that are infective on Rosaceae (Dryas, etc.), Coriariaceae (Coriaria), Datiscaceae (Datisca), and the genus Ceanothus (Rhamnaceae). Members of lineage 3 are the most promiscuous and are infective on Eleagnaceae, Rhamnaceae, Myricaceae, Gynmmostoma, and occasionally Alnus. The fourth Frankia lineage consists of the “atypical” strains which are unable to reinfect actinorhizal host plants or form ineffective root nodule structures that are unable to fix nitrogen. Our understanding of this genus has been greatly enhanced by the sequencing of several Frankia genomes from the different Frankia lineages [24-33]. Analysis of Frankia genomes has revealed new potential with respect to metabolic diversity, natural product biosynthesis, and stress tolerance, which may help aid the cosmopolitan nature of the actinorhizal symbiosis [31, 34].
In this chapter, we will describe what is known about the degradation properties of these bacteria.
Among bacteria with bioremediation potential, Frankia are unique in that these bacteria form a symbiosis with actinorhizal plants. The implications of this trait for bioremediation efforts have only recently been explored. In the context of bioremediation, the most extensively studied system is the Frankia–Alnus association. Diverse assemblages of free-living Frankia strains are present in soils with polyaromatic hydrocarbon (PAH) contamination [8-10, 15, 35-38]. These Frankia strains readily form symbioses with alders, resulting in greatly increased alder fitness in harsh environments. The Frankia–alder symbiosis also increases the mineralization of representative organic pollutants in oil-sands reclamation sites. The Frankia–alder symbiosis has been used in reclamation projects because of these traits [5, 8, 36-38]. Free-living Frankia also appears to be part of natural degradation communities. Specifically, Frankia has been found to be one of the most abundant genera in wastewater treatment communities [35]. Based on these findings, Frankia appears to be an underutilized tool in holistic remediation approaches.
Triazines are a class of herbicides composed of a heterocyclic six-membered ring with alternating carbon and nitrogen atoms joined by double bonds. These herbicides have been used extensively for control of broadleaf and grassy weeds in corn, sorghum, and sugarcane cultivation. Atrazine and simazine are the most ubiquitous members of the s-triazine family. Biodegradation of atrazine is a complex process and depends on the nature and amount of atrazine in soil or water [39-41]. There are four major steps in atrazine degradation: hydrolysis, dealkylation, deamination, and ring cleavage. For the hydrolysis step, an amidohydrolase enzyme (AtzA) cleaves the carbon-chlorine (C-Cl) bond and thus dechlorinates atrazine to hydroxylatrazine. This intermediate is dealkylated and deaminated at the ethyl and isopropyl groups by the amidohydrolase enzymes, AtzB and AtzC, to produce cyanuric acid. This product is converted to ammonia and carbon dioxide by the AtzD, AtzE, and AtzF enzymes [42-44].
In Frankia, the first two steps in atrazine degradation have been identified as well as the regulation of their gene expression [17]. The mineralization of atrazine to ammonia and carbon dioxide is generally initiated by hydrolytic dechlorination, catalyzed by the enzyme atrazine chlorohydrolase (AtzA). Alternatively, this reaction is catalyzed by another atrazine chlorohydrolase (TrzN), which is also able to use atrazine derivatives including desethyl-desisopropylatrazine as substrates. Analysis of the Frankia genomes identified candidate genes for the atrazine degradation pathway (Figure 1). The trzN gene was identified in Frankia alni ACN14a (FRAAL1474) and Frankia sp EuI1c (FraEuI1c_5874) genomes and its amidohydrolase gene product is predicted to remove chlorine from s-triazine compounds to produce hydroxyatrazine or ammeline from atrazine and desethyl desisopropyl atrazine, respectively. Furthermore, a putative atzB gene was also identified in both Frankia genomes (FRAAL1473 and FraEuI1c_5875) whose predicted gene product, adenosine aminohydrolase 3, is involved in the dealkylation reaction of the N-ethyl group from hydroxyatrazine transforming it into N-isopropylammelide. Physiological studies showed that Frankia ACN14a and EuI1c cultures are able to break down atrazine and desethyl-desisopropylatrazine producing the end products hydroxyatrazine and N-isopropylammelide. Although the enzymes were not purified, these data clearly showed metabolism of atrazine. Analysis of gene expression in Frankia ACN14a found that the two genes, trzN (FRAAL1474) and atzB (FRAAL1473) are under control of the atzR (FRAAL1471) gene, which encodes a predicted LysR-type transcriptional regulator.
Gene cluster organization in Frankia alni ACN14a for atrazine degradation. The cluster contains a putative trzN (FRAAL1474), putative atzB (FRAAL1473), and putative LysR-family transcriptional (atzR).
Bioinformatics analysis of the Frankia genomes revealed a potential full pathway for atrazine degradation in the Frankia sp EuI1c genome (Figure 2). The atzC (FraEuI1c_4724) gene, which encodes a putative amidhydrolase enzyme, was identified and is predicted to be involved in the dealkylation of the N-isopropyl group from atrazine to produce cyanuric acid. With other bacterial systems, cyanuric acid is hydrolyzed to ammonium and carbon dioxide via the atzDEF operon [43, 45]. In Frankia EuI1c, the atzD (FraEuI1c_3137) gene product is predicted to transform cyanuric acid into carboxybiuret, which spontaneously decarboxylates to biuret. Putative atzE (FraEuI1c_1007 and 1008), and atzF (FraEuI1c_3831) genes were also identified in the Frankia EuI1c genome and their gene products expected to complete s-triazine mineralization by converting biuret to allophanate and ammonia plus carbon dioxide. A trzR (FraEuI1c_3136) gene, which encodes a GntR family transcriptional regulator, is found before the atzD gene and is involved in the expression of that gene (Rehan unpublished).
The atrazine degradation steps in Frankia strains EuI1c and ACN14a include atrazine dechlorination and dealkylation and ring cleavage by TrzN, atzB, and atzD enzymes.
Biphenyls and polychlorinated biphenyls (PCBs) are some of the most recalcitrant xenobiotics found in the environment. The degree of chlorination differs greatly among the PCBs, ranging from 1 to 10, as does their position on the carbon atoms. Since the mid-1980s, the use of PCBs has been phased out in many countries. However, due to their toxicity, persistence in the environment, and potential carcinogenicity, they are still a major global environmental problem [46-48].
Bacteria degrade biphenyl and PCBs via the meta-cleavage pathway, which is encoded by the bph operon, and produces tricarboxylic acid and chlorobenzoate (CBA) as intermediates [47-50]. The first enzyme in this pathway is biphenyl dioxygenase, which is a multimeric complex consisting of the large α and small β subunits, and the ferredoxine and ferredoxine reductase subunits. The degradation process is initiated by biphenyl dioxygenase which incorporates two oxygen atoms at the 2 and 3 carbon positions of the aromatic ring (called 2,3-dioxygenation) to generate hydroxyl groups. For PCBs degradation, biphenyl dioxygenase catalyzes the initial 2,3-dioxygenation, and dihydrodiol dehydrogenase converts the product into 2,3-dihydroxybiphenyl. The enzyme, 2,3 dihydroxybiphenyl dioxygenase, cleaves the dihydroxylated ring to produce (chlorinated) 2-hydroxy-6-oxo-6-phenylhexa-2,4-dienoic acid (HOPDA). A hydrolase enzyme then hydrolyzes HOPDA to (chlorinated) benzoic acid and 2-hydroxypent a-2,4-dienoate.
At least four Frankia strains (ACN14a, CcI3, EUN1f, and EuI1c) are resistant to biphenyl and polychlorinated biphenyl (PCB) at concentrations up to 5mM [51, Swanson unpublished results]. Data mining for known organisms capable of biphenyl degradation [46, 52] and the availability of a Frankia genome database enabled the identification of genes potentially involved in biphenyl degradation in several of the Frankia strains listed above. Five genes were identified that encode enzymes involved in biphenyl degradation: the alpha and beta subunits of the aromatic-ring-hydroxylating dioxygenase, a Rieske (2Fe-2S) iron–sulfur domain protein, an alpha/beta hydrolase fold protein, and a short-chain dehydrogenase/reductase (SDR). These enzymes are putatively capable of oxidizing and hydroxylating benzene rings, and are also known as the upper meta-cleavage pathway. A lower pathway of aromatic ring degradation consisting of three genes (encoding the 2-hydroxypenta-2,4-dienoate hydratase; acylating acetaldehyde dehydrogenase; and 4-hydroxy-2-oxovalerate aldolase) is located downstream of this operon [53, Swanson and Tisa unpublished data]. Figure [3] shows the gene neighborhood of the Biphenyl degradation genes. These genes were also found in Frankia strain EUN1f and Dg1 genomes (Swanson and Tisa unpublished). Both the meta-cleavage upper and the lower pathways are commonly referred to as the bph operon in several other PCB-degrading bacteria. Rhodococcus RAH1, a species closely related to Frankia, utilizes bph genes homologous to those found in Frankia to metabolize PCBs as a sole carbon and energy source [54]. Since at least two genes (Aromatic-ring-hydroxylating dioxygenase, subunit alpha-like protein (FraEuI1c_4097) and short-chain dehydrogenase/reductase (FraEuI1c_4101) in the bph operon in Frankia are upregulated in the presence of biphenyl, it is likely that Frankia also uses the bph operon to metabolize biphenyl and PCBs (Rehan and Tisa unpublished)
The gene neighborhood of bph operon in Frankia EuI1c in comparison to Rhodococcus equi 103S and Photorhabdus luminescens laumondii TTO1 operon. (1) Aromatic-ring-hydroxylating dioxygenase, subunit alpha. (2) Rieske (2Fe-2S) iron–sulfur domain protein. (3) Aromatic-ring-hydroxylating dioxygenase, subunit beta. (4) Alpha/beta hydrolase fold protein. (5) Short-chain dehydrogenase/reductase SDR.
Phenol (or hydroxybenzene) consists of a benzene ring substituted with a hydroxyl group. Derivatives of this molecule are colloquially known as phenolic compounds. Phenolic compounds are ubiquitous chemicals with diverse properties and uses. The simplest phenolic compound, phenol, is widely used in oil and coal processing, tinctorial and metallurgic industries, and many other industrial applications. Phenol also enters the environment via vehicle exhaust and as the product of natural metabolic processes, and chlorophenols are widely used as biocides in agricultural applications [for a review see 55]. While anthropogenic phenolics are often hazardous, natural phenolic compounds are mostly harmless in the concentrations that are found in foods such as coffee and tea, and some are used as antibiotics [56, 57]. However, the toxicity of some phenolics, particularly phenol and chlorinated phenols, has prompted considerable research activity devoted to phenol remediation. Acute and chronic exposure to phenol and chlorophenol has serious health effects. Phenol and chlorophenol cause lipid peroxidation which ultimately leads to tissue necrosis, and liver and kidney damage [58]. Additionally, chlorophenol exposure is associated with elevated risks of cancer, immune deficiencies, and teratogenic effects [59-61].
One of the most promising techniques for removing anthropogenic phenolics from the environment is bioremediation. As was the case for many compounds, the degradation pathway for phenol was first elucidated in a Pseudomonas strain [62]. Most bacteria degrade phenolics using catechol catabolic enzymes, most importantly catechol-2,3-dioxygenase. Phenols are first hydroxylated to form catechol, and then catechol-2,3-dioxygenase cleaves the benzene ring at the meta position [62]. Therefore, the degradation pathway that begins with catechol-2,3-dioxygenase is called the meta pathway (Figure 4). While the meta pathway is most prevalent, degradation can also begin with cleavage at the para or ortho position using catechol-1,2-oxygenase [63-65]. After ring cleavage, 2-hydroxymuconic semialdehyde hydrolase catalyzes a decarboxylation reaction yielding 4-oxalocrotonate. 4-oxalocrotonate is hydrated by 2-oxopent-4-enoate hydratase to form 4-hydroxy-2-oxovalerate. 4-hydroxy-2-oxovalerate aldolase then splits 4-hydroxy-2-oxovalerate into pyruvate and acetaldehyde, which can then be incorporated into the central metabolic pathways [62].
General phenol degradation pathway.
Frankia spp. both produce and are affected by phenolic compounds. However, it is unclear whether Frankia may degrade phenol and other phenolic compounds. The response of Frankia to phenolics was first studied in the context of plant–microbe interactions. Despite apparent functional and morphological similarities between Frankia nodules and leguminous nodules, the molecular and physiological mechanisms that control nodulation are distinct. Therefore, the unique process of nodulation by Frankia is still an area of intense research. Alnus spp. (Alders) plants are a major host plant for Frankia, and also have unusually high levels of phenolics in their root exudates, which affect the growth of Frankia. Most Alnus phenolics tested inhibit Frankia growth to varying degrees [66, 67]. Specifically, benzoic acids are less inhibitory than cinnamic acids such as caffeinic acid. However, one plant phenolic, o-hydroxyphenylacetic acid, promoted Frankia growth, and both benzoic and cinnamic acids caused increased branching of Frankia hyphae. Low concentration plant phenolics also mediate a global shift in Frankia gene expression, while higher concentrations (above 30 mg L-1) simply inhibit biosynthesis [33]. Interestingly, Frankia also increases phenolic expression of their host plant, causing them to produce more phenol, flavonoids, and hydroxycinnamic acid [68].
Frankia may promote excretion of phenolics as a way to increase available nutrients. However, this explanation depends on Frankia having the ability to degrade phenolic compounds. While no study has demonstrated that Frankia degrades phenolic compounds, there is genetic evidence that this bacterium may have the ability to degrade phenolics. First, some Frankia strains have genes coding for the production of catechol and other phenolic compounds [34]. Because bacteria often salvage the biomolecules they produce, the presence of an anabolic pathway suggests that a catabolic pathway is also present [69]. Furthermore, multiple Frankia strains contain catechol-2, 3-dioxygenase, the most important enzyme in the phenol degradation pathway (Swanson and Tisa unpublished data) [64]. A closely related bacterium, Rhodococcus spp., uses the catechol-2,3-dioxygenase pathway to grow with phenol as its sole carbon source [70]. The same species is also able to break down the more recalcitrant pentachlorophenol via the para pathway [71]. This suggests that Frankia may break down phenol, a trait that could be applied in bioremediation efforts. Several Frankia strains are able to grow on phenol, quercetin, catechol, and other phenolic compounds (Furnholm, Greenleaf, and Tisa unpublished data), but the metabolism of their breakdown has not been studied.
Naphthalene is a ubiquitous polyaromatic hydrocarbon composed of two benzene rings joined at the 9 and 10 carbons (Figure 5). Naphthalene is produced by distilling and crystallizing coal tar, and also as by-product of fossil fuel combustion and cigarette smoke [72]. Naphthalene is used in a number of industrial applications including as feed stock for the production of plastics and resins, and as a component of creosote-based wood preservatives. Naphthalene is also used in tincture and leather tanning industries [72]. Unlike many organic pollutants, naphthalene does not bioaccumulate. Instead, naphthalene is metabolized and excreted in the urine of rats and humans [72, 73]. Nonetheless, naphthalene is a problematic pollutant with numerous toxic effects. Acute exposure to naphthalene causes hemolytic anemia, and liver and neurological damage [74]. Chronic naphthalene exposure is associated with elevated cancer risk [75, 76]. The toxicity of naphthalene and its prevalence as a pollutant has spurred research on remediation techniques, including bioremediation and biodegradation.
Structure of naphthalene.
The naphthalene biodegradation pathway was first studied in a strain of Pseudomonas which has two related naphthalene degradation pathways. The upper pathway catabolizes naphthalene to produce salicylate and a molecule of pyruvate [77]. The lower pathway breaks salicylate down into acetyl Co-A and pyruvate [78]. The first step of the upper pathway is catalyzed by four proteins: naphthalene dioxygenase reductase, naphthalene dioxygenase ferredoxin, and naphthalene dioxygenase Fe-S protein small and large subunits. This collection of enzymes oxidizes naphthalene to produce cis-naphthalene dihydrodiol, which is subsequently dehydrogenated by naphthalene cis-dihyrdodiol dehydrogenase to form 1,2-dihydroxynaphthalene. 1,2-dihydronaphthalene dioxygenase then produces 2-hydroxychromene-2-carboxylate which is then cleaved by 2-hydroxychromene-2-carboxylate dehydrogenase to form cis-o-hydroxybenzylpyruvate. 1,2-dihydroxybenzylpyruvate aldolase then splits cis-o-hydroxybenzylpyruvate producing pyruvate and salicylaldehyde. Finally, salicylaldehyde dehydrogenase carboxylates salicylaldehyde to form salicylate [77, 78].
In the lower pathway, salicylate hydroxylase hydroxylates salicylate to produce catechol. The remaining benzene ring is then cleaved by catechol-2,3-dioxygenase to produce 2-hydroxymuconic semialdehyde [78]. Hydroxymuconic semialdehyde dehydrogenase then produces 2-hydroxyhexa-2,4-diene-1,6-dioate which is subsequently isomerized by 4-oxalocrotmate isomerase to produce 2-oxohexa-3-ene-1,6-dioate. This is then transformed into 2-oxopent-4-enoate by 4-oxalocrotomate decarboxylase. 2-oxopent-4-enole hydratase produces 4-hydroxy-2-oxovalerate, which is subsequently split into acetaldehyde and pyruvate by 2-oxo-4-hydroxypentanoate aldolase. Finally, acetaldehyde dehydrogenase converts acetaldehyde into acetyl Co-A [78]. Both of these pathways are also found in Rhodococcus spp, a close relative of Frankia [79].
Not surprisingly, Frankia also metabolizes naphthalene as a sole carbon and energy source via a related pathway [18]. Specifically, Frankia uses the protocatechuate pathway to convert naphthalene or a naphthalene derivative into acetyl Co-A and succinyl Co-A (Figure 6) [18]. This finding confirms the role of Frankia in naphthalene degradation, which was suggested by earlier field studies [8-10, 37, 38]. In symbiosis with alders, Frankia increases polyaromatic hydrocarbon degradation in oil-sand tailings for the first 1.5 years [8, 10, 37]. However, after 2.5 years, alders without Frankia symbionts demonstrated naphthalene degradation equal to the degradation or Frankia-inoculated alders [8]. The Frankia-alder symbiosis thrives in PAH-contaminated areas [15]. Interestingly, alder plants found in these PAH-contaminated areas maintained a symbiosis with Frankia lineage III as opposed to the normal lineage I, suggesting that this pollutant affected nodulation and/or survival of the actinorhizal plants. Taken together, these findings indicate that Frankia could be a useful tool in naphthalene remediation.
Putative naphthalene degradation pathway in Frankia [18]. (Figure is recopied with permission from Canadian Journal of Microbiology.)
Under oxic conditions, microbial degradation of many aromatic compounds occurs through the catechol or protocatechuate branch of the ß-ketoadipate pathway via either ortho cleavage by catechol 1,2-dioxygenase and protocatechuate 3,4-dioxygenase or meta-cleavage by catechol-2,3-dioxygenase and protocatechuate-4,5-dioxygenase.
Besides the protochatechuate pathway found in Frankia QA3 [18], several other potential protocatechuate pathways have been identified from bioinformatics analysis of the available Frankia genomes. In Frankia EuI1c, a potential operon (FraEuI1c_2560 -to- FraEuI1c_2564) for a putative protocatechuate pathway was identified (Figure 7). This operon encodes the predicted gene products involved in the putative pathway including protocatechuate 3,4-dioxygenase alpha and beta subunits, fumarate lyase, 3-oxoadipate enol-lactonase, and 4-hydroxybenzoate 3-monooxygenase. These gene products are similar to the protocatechuate degradation pathway found in Rhodococcus opacus 1CP [80, 81]. These results suggest that Frankia may use the protocatechuate degradation pathway to degrade many aromatic ring compounds after their conversion to protocatechuate.
The proposed protocatechuate degradation pathway in Frankia strains EuI1c and EUN1f.
Petroleum-based energy and products are used extensively around the world. The pervasiveness of petroleum inevitably leads to serious environmental pollution. Petroleum is a complex mixture of hydrocarbons, cycloalkanes, aromatic hydrocarbons, and more complex chemicals like asphaltenes. These chemicals and their derivatives, which are termed petrogenic compounds, are released into the environment as a result of oil spills and combustion of petroleum-based products [82]. Oil spills are one of the most serious sources of petroleum pollution and devastate aquatic and marine environments. Ongoing research to identify new methods for petroleum remediation is important because oil spills and other types of petroleum-derived pollution continue to pose environmental health risks.
Hydrocarbon-degrading bacteria and fungi are widely distributed in marine and freshwater environments, as well as soil habitats [83, 84]. In Pseudomonas, the alkane hydroxylase (monooxygenase) system consists of three components: alkane hydroxylase (AlkB), rubredoxin, and rubredoxin reductase. This system is responsible for the first oxidation step in the utilization of n-alkanes [85]. Similar alkane hydroxylase systems have been found in a variety of alkane-degrading bacteria [86, 87]. Alcanivorax sp. strain 2B5 will degrade C13–C30 n-alkanes and branched alkanes (pristine and phytane) from crude oil as the sole carbon source via a novel alkane hydroxylase gene (alkB). Other Acinetobacter are able to use n-alkanes with chain length C10–C40 as a sole source of carbon. In addition, the presence of multiple alkane hydroxylases in two Rhodococcus strains were characterized and both organisms contained at least four alkane monooxygenase gene homologs (alkB1, alkB2, alkB3, and alkB4) [76, 88].
A bioinformatics approach was used to identify these potential hydrocarbon degradation pathways among the sequenced Frankia strains. Functionally analyzed genes for the known hydrocarbon degradation pathways [84, 88] were used to probe the Frankia genome database and identify potential pathways. Our preliminary results (Rehan unpublished data) revealed that the F. alni ACN14a genome possesses a putative alkane-1 monooxygenase (Alkane omega-hydroxylase) gene (FRAAL1986), which is one of the known enzymes involved in the breakdown of n-alkanes (Figure 8). Furthermore, a similar gene (Franean1_2192) was also found in the Frankia sp. EAN1pec genome. These bioinformatics results support the hypothesis that Frankia may be able to degrade oil-spill-derived hydrocarbons. However, these preliminary results need further study.
Potential alkane-1 monooxygenase identified in F. alni ACN14a.
Clearly, we have only begun to scratch the surface of the metabolism of Frankia and its biodegradative potential. These initial studies correlating metabolic capacity to gene function are the first step in exploiting the bacteria for their bioremediation ability. Further bioinformatics data mining are necessary to elucidate the unique metabolic potential of Frankia. However, these in silico studies require “wet lab” experiments to confirm these capabilities.
From limited field studies, actinorhizal nodule occupancy seems to be under control by environmental conditions. The presence of Frankia lineage III strains inside alder nodules found under PAH-stressed soils suggests that this lineage may have a greater metabolic potential. The larger genome size of this lineage compared to the other infective strains also supports this hypothesis. However, further experiments are required to confirm this postulate.
We thank Michele Greenleaf, Teal Furnholm, and Kaci B. Kus for their efforts on our degradation studies. Partial funding was provided by the New Hampshire Agricultural Experiment Station. This is scientific Contribution Number 2613. This work was supported by USDA National Institute of Food and Agriculture Hatch Project NH585. MR was supported by an Egyptian Channel Fellowship from The Egyptian Cultural Affairs and Missions Sectors.
Dry methane reforming (DMR) has drawn keen attention as viable CO2 utilization technology because it may have one of the greatest commercial potentials [1, 2].
Moreover, products are the main components of syngas (H2 and CO), which can be converted to the synthetic fuels as well as H2 carrier via well-established C1 chemistry. Conventionally, the H2/CO ratio from DMR is more suitable for Fischer-Tropsch synthesis than other methane reforming reactions [3, 4, 5]. Figure 1 shows the reaction enthalpy and Gibbs free energy of DMR (R1) with respect to temperature. According to the definition, reaction enthalpy (Eq. (1)) consists of two terms:
Energy diagram of DMR.
DMR is categorized as uphill (endothermic) reaction where energy input (ΔH) is indispensable in order to satisfy the conservation of energy. Moreover, the reaction does not occur spontaneously by the low-temperature thermal energy due to the large positive value of ΔG at low temperature. Figure 1 shows that at least 900 K is required to have a negative value of ΔG, and all energy is supplied via high-temperature thermal energy. Such high-temperature heat is supplied by the combustion of initial feed that produces CO2 as well as NOx. Net CO2 utilization is partly canceled unless combustion-generated CO2 is utilized which is economically quite difficult. Moreover, heat transfer from the combustion gas flowing outside of the reactor to the catalyst bed governs the overall material throughput which is known as a heat transfer-limiting regime. Because the heat transport property of a fixed bed reactor is poor, excessively high-temperature operation beyond thermodynamic limitation (i.e., 900 K) is necessary.
To overcome the aforementioned problem, low-temperature DMR is demanded, pursuing a new technology, and potential use of nonthermal plasma is highlighted. Assume DMR is operated at a lower temperature than the thermodynamic limitation as schematically depicted in Figure 1. A part of the energy is supplied by a low-temperature thermal energy (TΔS), while the rest of energy is supplied by the electricity (ΔG) under the nonthermal plasma environment so that TΔS + ΔG satisfies reaction enthalpy (ΔH). Electrical energy is used to accelerate electrons; subsequently, the electron energy is transferred to the molecules to initiate DMR at much lower temperature than thermal catalysis. Electronic collision process is independent of reaction temperature if gas density does not change significantly. Meanwhile, a part of the electrical energy is converted to heat: electrical energy consumed by nonthermal plasma (E) is depicted in the dashed line in Figure 1: inevitably, E is greater than ΔG at a fixed temperature. Although heat generated by nonthermal plasma is considered as energy loss (i.e., E−ΔG), both excited species and heat are utilized via endothermic DMR, which enables efficient use of electrical energy without heat transfer limitation: electrification of reforming reaction, or chemical processes in general, has the greatest advantage that the energy transfer and the control are independent of temperature gradient.
Dielectric barrier discharge (DBD) is the most successful atmospheric pressure nonthermal plasma sources in industry applications [6] and is used exclusively for this purpose. DBD is combined with a catalyst bed reactor and generated at atmospheric pressure [7]. DBD is characterized as a number of transient discharge channels known as streamers with nanosecond duration. Because the streamer has a nature of propagation along the interface between two adjacent dielectric materials, namely, the catalyst pellet and the gas interface, excited species produced by DBD is transferred to the catalyst surface efficiently. Moreover, the heat generated by DBD is transferred directly to the catalysts; overall energy transfer from nonthermal plasma to the catalyst bed is efficient. If the electricity is supplied from the renewable energy such as photovoltaics and wind turbines, low-emission DMR is possible with free of combustion. Moreover, nonthermal plasma-assisted C1 chemistry enables renewable-to-chemical energy conversion, which provides an alternative and viable solution for the efficient renewable energy storage and transportation pathways.
The aforementioned thermodynamic analysis (Figure 1) implies that the temperature-benign and low-emission chemical processes are possible with the appropriate combination of nonthermal plasma and the heterogeneous catalysts. Meanwhile, such hybrid system does not work at room temperature simply because the overall reaction rate is kinetically controlled at much low temperature: Nevertheless, we would like to highlight that nonthermal plasma technology solves many technological obstacles such as the elimination of combustion as well as heat transfer limitation. Moreover, low-temperature operation suppresses coke formation which is one of the big issues in DMR. In this book chapter, we focus on low-temperature DMR and compare thermal and plasma catalysis. Plasma catalysis of DMR was diagnosed by pulsed reaction spectrometry [8], and results were compared with thermal catalysis to highlight the benefit of DBD and catalyst combination. Subsequently, the interaction between DBD and catalyst pellets was discussed toward deeper insight into the mechanism. Finally, future prospects of plasma catalysis of DMR are provided.
Based on the location and number of plasma zone and catalyst bed, the combination of heterogeneous catalysts with plasma can be operated in three configurations: single-, two- and multistage, which are shown in Figure 2.
Schematic diagram of single-stage (a), two-stage (b), and multistage reactor (c). Catalyst is depicted as orange circle; plasma is depicted as purple “lightning” symbol.
In a single-stage reactor (Figure 2(a)), the catalyst is packed inside the plasma zone, where the interaction of plasma and catalysts occurs. Because thermal plasma (gas bulk temperature >3000°C [9]) could damage catalyst, single-stage reactor is, therefore, suitable for nonthermal plasma sources. The single-stage reactor is widely applied in CH4 reforming [10, 11, 12, 13, 14], direct conversion of CO2 [15, 16, 17, 18, 19], VOCs abatement [20, 21, 22], exhaust matter removal [23], formaldehyde removal [24], NOx synthesis [25], and ozone synthesis [6]. There are two significant merits of single-state reactor: (I) great flexibility exists in terms of electrode and reactor configurations that the reactor can be constructed using inexpensive materials such as glass and polymers and (II) reactive species, ions, electrons, etc. generated by nonthermal plasma could modify the gas composition, which affects the surface reactions with catalyst synergistically. However, the interaction between plasma and catalyst is complex when the catalyst is placed directly in the plasma zone. The synergy of plasma and catalyst will be discussed further in Section 4 based on the single-state reactor.
Figure 3 depicts a single-stage DBD reactor for CH4 reforming [26], mainly including a quartz tube reactor, high voltage (HV) electrode at the center, and ground electrode outside of the tube. Catalyst pellets are packed in the plasma zone between two electrodes, and both ends of the catalyst bed are fixed by metallic supports. The high voltage is applied between the HV centered electrode and ground electrode to generate dielectric barrier discharge over the pellet surface. Discharge power was measured by voltage-charge Lissajous analysis. The discharge gap, which is the distance between HV electrode and the ground electrode, is usually less than 10 mm [27]. Catalyst temperature is controlled by a furnace. The temperature distribution of the catalyst bed is measured by thermography through the observation window. Figure 3(c)–(e) shows an overview of the catalyst bed, DBD generated in the catalyst bed, and the temperature distribution during reforming reaction. The catalyst bed temperature was clearly decreased because of the endothermic nature of DMR. In addition, gas temperature was estimated by optical emission spectroscopy (OES) of CO(B-A) transition [28], showing that catalyst temperature and gas temperature matched within a measurement error.
Single-stage DBD reactor system for DMR: (a) overview of the reactor system, (b) cross-sectional view, (c) overview of the catalyst bed, (d) DBD generated in the catalyst bed, and (e) temperature distribution during reforming reaction.
In the two-stage reactor, the catalyst is located at the downstream of plasma (Figure 2(b)). The gas is first addressed by the plasma and subsequently interacts with the catalyst [29]. Due to the separation of plasma and catalyst, both thermal and nonthermal plasma can be utilized. Because excited species generated in plasma have very short lifetimes, plasma mainly plays the role to preconvert the gas composition and then feed it into the catalyst reactor, e.g., in NOx removal process, due to the pretreatment of plasma, NO and NO2 were coexisted, which enhanced the following selective catalytic reduction in catalyst bed [30]; the other example is the benzene removal process where ozone (O3) was formed from background O2 by plasma, which promoted the decomposition of benzene in the next stage [31]. However, compared with the single-stage reactor, application of a two-stage reactor is limited in plasma catalysis and shows a lower performance for a given catalyst [32, 33, 34, 35, 36, 37].
The multistage reactor can be described as a combination of more than one single-stage bed/reactor (Figure 2(c)). The multistage reactor gives a more flexible option in the industrialization of the plasma catalysis, attributing to the combination of catalysts with a different function for the expected reaction [38]. Chavade et al. [39] used a four-stage plasma and catalytic reactor system for oxidation of benzene. The results showed that the increase in stage number enhanced benzene conversion and CO2 selectivity. The same result can be found in biogas reforming process using a multistage gliding arc discharge system without catalyst [40]. Harling et al. [41] developed a three-stage reactor for VOCs removal. The combination of plasma and catalyst in series could significantly improve the efficiency of VOCs decomposition. At the same time, the formation of by-product such as NOx was suppressed.
The pulsed reaction spectrometry using DBD with Ni/Al2O3 catalysts was investigated to develop a reforming diagnostic method [10]. Pulsed reforming enables the transient analyses of both CH4/CO2 consumption and H2 and CO generation. Furthermore, carbon formation was analyzed quantitatively without serious catalyst deactivation. The varied CH4/CO2 ratios between 0.5 and 1.5 were investigated at a fixed catalyst temperature near 600°C. The CH4/CO2 ratio was initially set to 0.5, and then the CH4/CO2 ratio was incremented stepwise until CH4/CO2 = 1.5, consecutively, while total flow rate was fixed at 1000 cm3/min. De-coking process (R2) was followed up after every pulsed reaction. System pressure was kept at 5 kPa during the reforming process. Discharge power was 85−93 W where specific energy input was ca. 1.2 eV/molecule. Commercially available catalyst pellets (11 wt% Ni-La/Al2O3, Raschig ring type: 3 mm) was packed for 40 mm length (total weight ca. 12 g; Ni 1.36 g; La 0.35 g). Figure 4 provides an overview of gas component changes in the entire hybrid reforming.
Overview of the entire pulsed hybrid DMR.
Reactant conversion and product yields are shown in Figure 5. The definition for conversion and yield were provided in Ref. [8]. CH4 conversion and H2 yield were monotonically increased with the CH4/CO2 ratio. There are two simultaneous routes for CH4 conversion as shown in Figure 6. Route (I) is a reforming path: CH4 is chemisorbed on metallic sites (adsorbed species are denoted by * in reaction). The adsorbed CH4 fragments (CHx*) is oxidized by CO2* to form CHxO* before complete dehydrogenations to C* occurs. In route (II), CH4 almost irreversibly dehydrogenates toward carbon atom, and then C*-rich layer is oxidized slowly by CO2* (R2), which can be evidenced in the de-coking process in Figure 4. When the CH4/CO2 ratio exceeded 1.0, CH4 prefers to dehydrogenate to solid carbon through route (II) due to the low proportion of CO2. Subsequently, a nonnegligible amount of solid carbon is produced, and CO2 conversion and CO yield turned to proportionally decrease.
Effects of CH4/CO2 ratio on DMR at ca. 600°C: (a) CH4 conversion, (b) CO2 conversion, (c) H2 yield, and (d) CO yield.
Two simultaneous routes for CH4 conversion.
Compared with thermal reforming, both CH4 conversion and H2 yield were clearly promoted in hybrid reforming (Figure 5), and the main pathway of CH4 conversion and H2 yield could be simply described as CH4 dehydrogenation (R3). It is proposed that CH4 dehydrogenation was enhanced by the synergistic effect of DBD and catalyst. Molecular beam study revealed that dissociative chemisorption of CH4 on the metal surface was enhanced by vibrational excitation [42]. The numerical simulation of one-dimensional streamer propagation demonstrated that the vibrationally excited CH4 is the most abundant and long-lived species generated by low-energy electron impact [43]. The reaction mechanism of plasma-enabled catalysis could be explained by the Langmuir-Hinshelwood (LH) reaction scheme. The analysis of overall activation energy is expected to understand the contribution of plasma-generated reactive species.
The CO2 conversion and CO yield were promoted in hybrid reforming compared to thermal reforming (Figure 5). H2O was simultaneously produced as a by-product by reverse water gas shift (RWGS) reaction (R4). Reactivity of plasma-activated H2O was confirmed by Arrhenius plot analysis where reaction order for H2O was doubled by DBD [44]. Plasma-activated H2O promotes reaction with adsorbed carbon; it creates additional pathways (R5) to syngas (H2 and CO). The CO2 conversion and CO yield were promoted in the hybrid reforming, illustrating that the reverse-Boudouard reaction (R2) was enhanced by DBD. The reaction between plasma-activated CO2 and adsorbed carbon increases CO yield. The same result was obtained in the de-coking period [10]. Although excessive production of carbon is detrimental for catalyst activity and lifetime, the presence of adsorbed carbon creates key pathways for emerging plasma-induced synergistic effect. Consequently, plasma-activated CO2 and H2O would promote surface reaction and increase CO and H2 yield. Figure 5 clearly shows that the slope of each line increased in hybrid reforming compared with thermal reforming, attributing to the nonthermal plasma-excited species. The increase of slope could be further explained by the promoted overall reaction order, which plays the key role in the estimation of the rate-determining step [44].
Synergism in plasma catalysis in the single-stage reactor is not fully understood due to the complex interaction between the various plasma-catalyst interaction processes [45, 46, 47, 48, 49]. Kim et al. [27] discussed the criteria for interaction between nonthermal plasma and the porous catalyst. The chemical species in nonthermal plasma are highly reactive; the lifetime is very short, e.g., O (1D), 10 ns; O (3P), 50 μs; and OH, 100 μs: their one-dimensional diffusion length is limited from 0.7 to 65 μm. Plasma generated species within diffusion length from the external surface of pellets would contribute to the plasma-induced reaction pathways. The interdependence of plasma and catalyst can be discussed as two aspects: catalyst affects plasma, and plasma affects catalyst.
With the packed catalyst in the plasma zone, gaseous species adsorbed on the catalyst surface increase the concentration of surface species. In addition, the electric field is enhanced near the catalyst surface due to catalyst nanofeatures [50, 51]. Moreover, the packed catalyst also enables the discharge type change, as well as microdischarge generation.
Without packed materials, discharge mode is the “free-standing” filamentary discharge propagating across the gas gap (Figure 7(a)). With a packed catalyst, the surface streamer is propagated with the close contact with the catalyst surface, and intensive partial discharges occur between the contact area of catalysts [52, 53]. The time-resolved visualization of surface streamer propagation and partial discharge were detected by Kim et al. [54] with an intensified charge-coupled device (ICCD) camera. Figure 7(b) shows time-resolved images of surface streamers (i.e., primary surface streamer and secondary surface streamer) propagating on the surface. Enhanced catalytic performance in the presence of a catalyst is closely linked with the propagation of surface streamers [55]. Moreover, with packed catalyst, microdischarge is generated inside catalyst pores (when the pore sizes >10 μm) [56, 57]. Zhang et al. [56] investigated microdischarge formation inside catalyst pores by a two-dimensional fluid model in the μm range (Figure 7(c)), indicating that the plasma species can be formed inside pores of structured catalysts in the μm range and affect the plasma catalytic process.
(a) Filamentary discharge without catalyst pellet, (b) time-resolved images of surface streamers propagating on the surface of γ-Al2O3 (reproduced with permission from ref 54. Copyright 2016 IOP Publishing), and (c) distributions of the electron density and total ion density with a 100 μm pore (reproduced with permission from ref 56. Copyright 2016 Elsevier).
The plasma also affects the catalyst properties, which are summarized as the following aspects:
Modification of physicochemical properties of the catalyst by plasma, which is widely used in catalyst preparation processes [58]. With plasma preparation, the catalyst obtains a higher adsorption capacity [59], higher surface area, and higher dispersion of the catalyst material [60, 61, 62], leading to a plasma-enhanced reactivity.
It is possible that plasma makes changes in the surface process with the catalyst. As for the CH4 reforming process, the deposited carbon from Ni catalysts can be removed effectively by plasma-excited CO2 and H2O [8]. The other example of the synergism is NH3 decomposition for the application of fuel cell, where NH3 conversion reached 99.9% when combining plasma and catalyst, although the conversion was less than 10% in the case of either plasma or catalyst only reaction [63].
Based on Arrhenius plot analysis, plasma can decrease the activation barriers (overall activation energy), attributed to the vibrational excitation, which is schematically depicted in Figure 8. The net activation barrier will be
The excited species or dissociated species might create other pathways with the presence of catalyst, e.g., during the CO2 plasma oxidation process, plasma-enhanced CO2 oxidized Ni/Al2O3 catalyst to form a NiO layer, which could drive an oxidation–reduction cycle in dry methane reforming reaction. The same NiO layer was found when specific energy input (SEI) was sufficiently high: the O2 that dissociated from CO2 plays the key role in the oxidation process. The details will be interpreted in Section 5.2.
Reduction of the overall activation energy by vibrational excitation of the reactants. (A) Adiabatic barrier crossing case and (B) nonadiabatic barrier crossing case. Reprinted with permission from ref 64. Copyright 2004 AAAS.
Coke formation behavior was studied as a reaction footprint to track reaction pathways induced by DBD [26]. Coke morphology and their distribution over the 3 mm spherical Ni/Al2O3 catalyst pellets were obtained after 60 min DMR. Figure 9 shows cross-sectional carbon distribution, where (a)–(c) and (d)–(f) correspond to plasma catalysis and thermal catalysis in low, middle, and high temperatures. For the thermal catalysis with the temperature at 465°C, carbon deposition over the entire cross-section was obvious. With the temperature increased, coke was decreased and finally became nondetectable at ca. 620°C. At low temperature, plasma catalysis suppressed the coke formation significantly over the entire cross-section.
Carbon distribution over the 3 mm spherical pellet cross-section after 60 min reforming: plasma catalysis (a)–(c) and thermal catalysis (d)–(f), respectively. Reprinted with permission from ref 26. Copyright 2018 IOP Publishing.
By the analysis of scanning electron micrographs (SEM) and Raman spectrum, fine carbon filaments were detected on the external pellet surface in plasma catalysis [26]. In contrast, thick fibrous carbon deposition was observed on the external surface in thermal catalysis, as well as in the internal pores in both thermal and plasma catalyses. The CH4 dehydrogenation on catalyst is enhanced by nonthermal plasma, contributing to the generation of highly filamentous and amorphous carbon. Such nonthermal plasma-enhanced reaction has been demonstrated by carbon nanotube growth [66] and plasma-enabled steam methane reforming [67]. The fine amorphous carbon filaments, deposited in the external surface of catalyst, prove that the interaction of DBD occurs mainly in the external surface. Consequently, DBD generation and plasma-excited species diffusion are inhibited in the internal pores of catalyst.
The nickel (Ni) of Ni/Al2O3 catalyst was oxidized slightly by CO2 in the thermal catalysis [68, 69] . However, the significant Ni oxidation by CO2(R6) was demonstrated when the catalyst were packed in nonthermal plasma zone. In this case, Ni uptakes surface oxygen beyond the adsorption/desorption equilibrium (i.e., Langmuir isotherm) to form NiO, which further promotes CH4 dehydrogenation without solid carbon deposition (R7):
The specific energy input (SEI) is a critical operational parameter in plasma-enabled CO2 treatment process due to the fact that dominant reaction pathway shifts dramatically with SEI:
SEI expresses energy consumption by discharging per unit volume of the feed gas, which could be further interpreted as average electrical energy (eV) per molecule. In Eq. (2), C is the conversion factor of the unit [10]. Two contrasting conditions are demonstrated in plasma-enabled CO2 oxidation: one is designated as the direct oxidation route; with a small SEI, CO2 dissociation to CO and 0.5 O2(R9) is negligible, and then the plasma-excited CO2 dominates the oxidation process (R8). The other is the indirect oxidation route where O2 provides an additional oxidation pathway; with a large SEI, CO2 is dissociated into CO and O2(R9) without heterogeneous catalysts by electron impact [70, 71], followed by Ni oxidation by O2(R10).
The plasma-enhanced direct oxidation route (R8) is further investigated because the plasma-enabled synergistic effect was demonstrated distinctly without O2 [8, 10, 26]. Ni oxidation behavior without O2 was studied with SEI = 0.46 eV/molecule. The CO2 conversion is far below 1% when the SEI was smaller than 0.5 eV/molecule [72, 73]; in the plasma and thermal oxidation, the CO2 flow rate, catalyst temperature, and the oxidation time were controlled as 1000 cm3/min, 600°C, and 70 min, respectively.
After DBD-enhanced oxidation and thermal oxidation, the formation of NiO and its distribution over the cross-section of 3 mm spherical pellet were investigated by Raman spectroscopy and optical microscope. Results showed that the NiO layer was recognized clearly with the thickness of ca. 20 μm. In contrast, the NiO layer was not identified after thermal oxidation. We should point out that the plasma-excited CO2 has a strong oxidation capability of Ni catalyst. In addition, the effect of DBD is inhibited in the internal pores beyond 20 μm from the pellet surface.
In the thermal oxidation, CO2 is most likely adsorbed at the perimeter between Ni nanocrystals and Al2O3 interfaces [74, 75, 76] (Figure 10(a)). Subsequently, the adsorbed CO2 oxidize Ni to NiO near the perimeter. It is clear that the reaction sites for thermal oxidation are limited in the perimeter. The Ni oxidation reaction terminates after the reaction sites are fully oxidized by adsorbed CO2. In plasma-enhanced oxidation reaction, CO2 is firstly excited by electron impact. The vibrationally excited CO2 plays the key role to enhance adsorption process and subsequent oxidation reaction of Ni catalyst, leading to an extensive Ni nanoparticle oxidation, which occurs not only in the perimeter but also in the terrace, step, and kink (Figure 10(b)). Figure 10(c) and (d) show hemispherical catalyst pellets after thermal and plasma oxidation. After thermal oxidation, the external surface and cross-section of catalyst pellets remained black. In contrast, after plasma oxidation, the external surface was oxidized and has showed whitish color change (oxidized stage); in the meantime, the cross-section of the hemispherical pellet has been kept black (unoxidized stage).
Ni oxidation pathways: (a) thermal oxidation including CO2 adsorption near the perimeter of Ni catalysts and (b) plasma-enhanced oxidation. Hemispherical catalyst pellets: (c) thermal oxidation and (d) plasma oxidation.
The vibrationally excited CO2 by DBD would induce Ni oxidation to form the oxygen-containing active species (i.e., NiO) rather than simple adsorption, leading to oxygen-rich surface beyond Langmuir isotherm. Incoming plasma-excited CO2 would carry a few eV internal energy due to the gas phase vibration-to-vibration energy transfer [77, 78], which is the main source of energy for NiO formation. Plasma-excited CO2 could promote the adsorption flux; however, the adsorbed CO2 is finally desorbed by the equilibrium limitation unless it forms NiO. In addition, the plasma-induced nonthermal heating mechanism plays another key role in the enhancement of Ni oxidation. Charge recombination and association of radicals can release energy corresponding to 1–10 eV/molecule on catalyst surface. When this excess energy is directly transferred to the adsorbed CO2, Ni oxidation may be enhanced without increasing macroscopic catalyst temperature. This reaction scheme is explained by nonthermal plasma-mediated Eley-Rideal mechanism, rather than precursor-type adsorption enhancement.
In the DMR process, the oxygen-rich surface (NiO layer) has a capability of oxidizing a large flux of ground-state CH4 efficiently. Consequently, CH4 is not necessarily preexcited. CH4 is almost fully reacted in the NiO layer (20 μm thickness) to inhibit the coke deposition toward the internal pores [26]. However, in the thermal catalysis, NiO is generated in a negligible amount. The ground-state CH4 can diffuse into internal pores and deposit coke as previously confirmed [26]. Formation of NiO shell (Figure 10(d)) and the coke formation behavior (Figure 9) are well correlated in plasma catalysis as further discussed in the next section.
Although the synergies of plasma and catalyst have been summarized in Section 4, the interaction between DBD and catalyst pores will be further discussed in this section based on the carbon formation and oxidation behavior, as well as DBD-enhanced DMR. For the plasma catalysis, carbon deposition in the internal pores could be remarkably prevented, and fine amorphous carbon filaments were deposited only on the external surface of pellets. A similar trend was observed when NiO was formed in the limited region over the external surface (20 μm depth) only when DBD was superimposed. The results of coke formation behavior and oxidation behavior of Ni-based catalyst in plasma catalysis evidence that the interaction of DBD and catalyst occurs at the external surface of the pellets and the effected thickness is ca. 20 μm. Neither generation of DBD nor diffusion of plasma-generated reactive species in the internal pores is possible. Although DBD and pellet interaction is limited in the external surface, conversion of CH4 and CO2 was promoted clearly compared with thermal catalysis: this is the clear evidence of reaction enhancement by DBD.
For DBD, due to the enhanced physical interaction between propagating streamers and catalysts, plasma and catalyst contact area, as well as the streamer propagation from one pellet to the other, are promoted significantly. Nevertheless, electron density in a narrow filamentary channel is of the order of 1014 cm−3 [6, 79, 80]; in contrast, molecule density at standard condition is approximately 1019 cm−3, indicating that a major part of the gas stream is neither ionized nor excited. Consequently, the extremely low proportion of ionized and excited species is inadequate to explain the net increase of CH4 and CO2 conversion and selectivity change by DBD. However, if reactive species are fixed and accumulated on the surface of the catalyst, the gross conversion of materials will be promoted. For this reason, the hetero-phase interface between DBD and the external pellet surface provides the most important reaction sites. In Section 5.2, nonthermal plasma oxidation of Ni to NiO creates a critically important step for plasma-enabled synergistic effect.
As Section 4.1 mentioned, gas breakdown is hard to occur in a pore smaller than 10 μm. For the pores catalyst with a pore size less than 2 nm, standard Paschen-type gas breakdown is impossible. To sum up, the external surface of pellet plays the key role for the DBD and catalyst interaction; however, the internal pores play a minor role.
The synergistic effect induced by DBD was clearly observed both in the CH4 and CO2 conversion and in the syngas yield. CH4 dehydrogenation was enhanced by the synergistic effect of DBD and catalyst. Plasma-activated CO2 and H2O would promote surface reaction and increase CO and H2 yield. The analysis of overall activation energy is expected to understand the contribution of plasma-generated reactive species.
In plasma catalysis, the fine amorphous carbon filaments, deposited in the external surface of catalyst, prove that the interaction of DBD occurs mainly in the external surface. The DBD generation and plasma-excited species diffusion are inhibited in the internal pores of the catalyst. Moreover, although the interaction between plasma and catalyst is limited in the external surface, the coke deposition was inhibited significantly in the internal pores by DBD, which is the clear evidence of reaction enhancement by DBD.
Oxidation behavior of Ni-based catalyst in nonthermal plasma-enabled catalysis showed that the NiO layer was generated in the external surface with the thickness of ca. 20 μm during plasma oxidation. In the internal pores, Ni oxidation is inhibited due to the negligible interaction with DBD. Contributing to the NiO layer, the surface of catalyst uptakes more oxygen beyond thermal equilibrium, which is known as Langmuir isotherm, creating a new reaction pathway via NiO. In the plasma catalysis of DMR, NiO drives the oxidation-reduction cycle, which promotes CH4 dehydrogenation on the surface. Consequently, carbon deposition is suppressed effectively.
For further improvement of plasma-enhanced DMR, the following issues should be investigated:
The effect of radical injection on reaction enhancement should be kinetically analyzed by the Arrhenius plot method, and the analysis of the overall activation energy is expected to understand the contribution of plasma-generated reactive species.
Exploring new types of catalysts, dedicated to plasma catalysis, is an important subject of research. We have demonstrated that the interaction of DBD and catalyst occurs only at the external surface of the pellets, and the effected thickness is ca. 20 μm, which means a majority of the active sites in pores of catalyst do not interact with any excited species. New catalyst preparation method such as catalytic functionalization of reactor wall and catalyst coating for the reactor may be beneficial to strengthen the synergistic effect of nonthermal plasma and catalytically functionalized surface.
The catalyst activity of partially oxidized catalyst and the nonthermal plasma heating mechanism have not been demonstrated experimentally yet; moreover, diagnosis of intermediate species on the surface, created by plasma-derived species, as well as their reaction dynamics are expected to be investigated for deep insight in plasma catalysis.
Although the plasma-induced energy transfer mechanism is commonly accepted in particle growth, it has yet to be investigated within the scope of plasma catalysis. Deep understanding of highly transient and nonequilibrium energy transfer via excited molecules, without macroscopic temperature change, need to be studied.
The individual contribution of radical injection and heat generation, as well as combination of those, must be understood. The gap between macroscopic and microscopic understanding, including various time scales covering nanoseconds to the millisecond, should be bridged by consistent manner.
Authors are listed below with their open access chapters linked via author name:
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\\n\\nJim Van Os 2015-18
\\n\\nLong Wang 2017, 2018
\\n\\nFei Wei 2016-18
\\n\\nIoannis Xenarios 2017, 2018
\\n\\nQi Xie 2016-18
\\n\\nXin-She Yang 2017, 2018
\\n\\nYulong Yin 2015, 2017, 2018
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\n\n\n\n\n\n\n\n\n\nJocelyn Chanussot (chapter to be published soon...)
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