scieee Open visual document viewer

Is the Ocean of Enceladus in a Primitive Evolutionary Stage?

Villavicencio Valero, K.; Ramírez Juidias, Emilio; Ávila Bosch, A.

Abstract

Enceladus has a subsurface ocean in the South Pole that has been inferred due to the presence of water vapor and other molecules like molecular hydrogen and ammonia detected by the Cassini mission from the ejection of material through the plumes in that region. The chemical composition of this ocean could give some information about the evolutionary stage of the icy moon if its components are found to be similar with the aqueous chemistry of the primitive oceans on Earth during glacial periods. Here we present a comparative geochemical analysis between the ocean of Enceladus and the aqueous composition of the oceans on Earth during the Snowball Event, in order to figure out if there are similar species, how the interaction of the metabolic processes between them works and if, in the future, those molecules could evolve making possible the emergence of life.

Full text

Selec ion o ou books indexed in he Book Ci a ion Index in Web o Science™ Co e Collec ion (BKCI) In e es ed in publishing wi h us? Con ac book.depa men @in echopen.com Numbe s displayed abo e a e based on la es da a collec ed. Fo mo e in o ma ion isi www.in echopen.com Open access books a ailable Coun ies deli e ed o Con ibu o s om op 500 uni e si ies In e na ional au ho s and edi o s Ou au ho s a e among he mos ci ed scien is s Downloads We a e In echOpen, he wo ld’s leading publishe o Open Access booksBuil by scien is s, o scien is s 12.2% 183,000 195M TOP 1% 154 6,800 Chap e Is he Ocean o Enceladus in a P imi i e E olu iona y S age? Ka he ine Villa icencio Vale o, Emilio Ramí ez Juidías and Aina À ila Bosch Abs ac Enceladus has a subsu ace ocean in he Sou h Pole ha has been in e ed due o he p esence o wa e apo and o he molecules like molecula hyd ogen and ammo- nia de ec ed by he Cassini mission om he ejec ion o ma e ial h ough he plumes in ha egion. The chemical composi ion o his ocean could gi e some in o ma ion abou he e olu iona y s age o he icy moon i i s componen s a e ound o be simila wi h he aqueous chemis y o he p imi i e oceans on Ea h du ing glacial pe iods. He e we p esen a compa a i e geochemical analysis be ween he ocean o Enceladus and he aqueous composi ion o he oceans on Ea h du ing he Snowball E en , in o de o igu e ou i he e a e simila species, how he in e ac ion o he me abolic p ocesses be ween hem wo ks and i , in he u u e, hose molecules could e ol e making possible he eme gence o li e. Keywo ds: ocean, snowball e en , aqueous chemis y, species, li e 1. In oduc ion Enceladus, one o he moons o Sa u n, p esen s a global ocean benea h he ice shell [1]. The exis ence o ha ocean was sugges ed because o he wa e apo de ec ed by he Cassini mission, h ough he ejec ion o ma e ial om he plumes loca ed in he sou h pole [2, 3]. The expulsion o ma e ial om he wa e plumes could be ela ed o hyd o he mal ac i i y [4], whe e ice pa icles a e hea ed due o he idal de o ma ion [5] and expelled o he su ace. E idence ha hose pa icles a e associ- a ed o hyd o he mal ac i i y a e he silica e sal s esidues ound a he E- ing [6], and he small size o he nanopa icles o ha ing. Bo h cha ac e is ics indica e ha he possible liquid wa e wi hin he ocean laye was p e iously in con ac wi h a ho silica e en i onmen [7]. The Ion and Neu al Mass Spec ome e (INMS) ins umen on boa d o he Cassini mission also de ec ed ammonia and some aces o o ganic molecules like benzene [8]. Ammonia is one mo e clue o he p esence o liquid wa e . Residuals om ammonia a e ni ogen-bea ing and oxygen-bea ing molecules ha , in combina- ion, could con e in o amino acids like i happens on Ea h [9]. O he de ec ed species we e H2O and CO2[10]. The me abolic in e ac ion be ween hese la e wo 1 species, h ough me hanogenesis, can o m me hane. I was also de ec ed molecula hyd ogenH2by he Cassini spacec a [11]. The e we e also ound some species wi h compounds o ca bon, ni ogen, oxygen and sul u [8, 12, 13]. The in e ac ion be ween molecula hyd ogen and some ca bona es wi hin he ocean p oduce a chem- ical ins abili y ha cons i u es an ene gy sou ce ha may suppo li e [11]. The Cosmic Dus Analyze (CDA) aboa d o he Cassini mission de ec ed wa e ice, o ganic molecule, and siliceous ma e ial [14]. The e we e also de ec ed concen- a ions o Na, and some sodium sal s like NaCl, NaHCO3and Na2CO3indica ing he p esence o liquid wa e [15, 16]. To main ain his liquid wa e in o he global ocean, he idal dissipa ion could be conside ed as an ene gy sou ce ha come om inside Enceladus [17]. Tidal hea ing also ac s in he solid co e p o oking high empe a u es in o he hyd o he mal ac i i y [18]. The hyd o he mal ac i i y c ea es con ec ion columns ha p oduce a dynamic mo emen in he ocean anspo ing he hea in o he ice shell om he co e [19]. The dissipa ion o he hea linked o he gas a ios p esen in he wa e plume could de e mine he s a e o he hyd o he mal ac i i y [7]. Acco ding o Woods [20], a hyd o he mal sou ce o gas could explain he dis ibu ion o hyd ogen in he wa e plume. In his sense, i mus be emphasized ha he abundance o hyd ogen de ec ed is simila o some aces o ola ile compounds like ca bon dioxide, me hane, and ammonia [8]. Labo a o y simula ions [7] sugges ed ha , in Enceladus, he molecula hyd ogen is a p oduc o in e nal eac ions. E idence o he in e nal p oduc- iono molecula hyd ogenis hehigh a ioo H 2=H2O which canno come om a gas apped in he ocean, because o i s high concen a ion, and i canno also come om he emnan s o a o ma ion en i onmen , due o he low a io o He=H2[21]. The geochemical sys em o he ocean o Enceladus could be composed mainly byNa2O -HCl-CO2-H2O. The concen a ion o CO2in he plumes is assumed o be he same ha may be ound in he dissol ed molecules o he subsu ace ocean. I s p esence sugges s a basic pH o he ocean o Enceladus. The es ima ion o his pH is based on he s udy o he he modynamic equilib ium, conside ing he empe a u e close o 0°C, he p essu e a 1 ba , he ca bon dioxide ac i i y, he chlo ide concen- a ion, and he dissol ed ino ganic ca bon HCO3=CO32. Acco ding o Glein e al. [22], he pH is 12.15 1.15. Ca bona es and bica bona es ions CO32=NaCO3a e also p esen in he ocean [23] and could come om soluble ca bona e mine als, o med h ough he eac ion o a apped CO2and silica mine als du ing wa e - ock di e en ia ion. I he ocks o he ocean eac wi hCO2, i is easible he ca bona ion p ocess o high wa e - ock a ios [24]. The me al concen a ion in he seawa e is o med by phyllosilica es and hyd oxide mine als, ha need an acid in o de o hyd olyze and being inco po a ed in o ca bona e mine als [23]. CO2=H2O a io in he plume o Enceladus is simila o he a io ound in he seawa e on Ea h [23], whe e heCO2ac i i y is con olled by al e a ion o mine als, assuming ha wa e - ock in e ac ions a e he main d i ing o ce o he pH as well as he composi ion o he ocean. On Enceladus, he se pen iniza ion is assumed o be he esul o mine als al e a ion [25], h ough a hyd olysis o p ima y mine als con aining i on and magnesium which p oduc is he hyd ogen. This p ocess is usually associa ed wi h ul ama ic ocks (<45% o SiO2and high Mg - Fe con en ), which eac ion ha akes place is he oxida ion by wa e o Fe(II) and Fe(III) in mine als such as oli ine and py oxene. The p oduc o his eac ion is he molecula hyd ogen. The p esence o hyd ogen can o m linea chains o hyd oca bons like me hane CH4 om he chemical eac ion be ween CO2and H2. This me hane could be p esen 2 As onomy and Plane a y Science - F om C yo olcanism o Black Holes and Galac ic E olu ion in low concen a ions in o he ocean o Enceladus [26]. E idence o his, i is he o ma ion o cla h a es ha a e able o ap ce ain molecules, which hen, would ise o he su ace and e en ually dissocia e and en ich he plume wi h me hane. Me hanol was also de ec ed by he Cassini mission; i is possible ha his compound has a biological o igin [27]. I was ound ha he CH3OH=H2O a io has ce ain co ela ion wi h bio ic ac i i y a ound he hyd o he mal en s. The concen a ion o me hanol de ec ed in he a mosphe e is high, which gi es a clue abou ha his specie is o med benea h he ice shell be o e being expelled in o he a mosphe e. These o ganic com- pounds de ec ed could be conside ed as a building block o li e o e en by-p oduc s o li e [28]. On Ea h, he i s signs o li e came om he A chean oceans whe e he oxida i e eac ions we e a p oduc o he in e ac ion be ween molybdenum and henium [29]. The e we e only aces o oxygen be o e he G ea Oxygen E en bu hen, a e i , he pho osyn he ic ac i i y led o an inc emen o his elemen [30]. The e olu ion o oxygen in he a mosphe e and oceans wen h ough i e s ages [31]. Du ing he C yogenian age, he a mosphe e and he shallow oceans had an inc ease o oxygen. The oxygen concen a ion was s agnan in ha e a, and subsequen ly i had an inc e- men ha con inued a e he nex million yea s and migh ha e culmina ed a ound he Ca boni e ous age. Du ing glacial pe iods, he concen a ion o CO2in he a mo- sphe e d opped and, be o e he eme gence o pho osyn he ic li e, he ca bon dioxide was mo e abundan in he a mosphe e han nowadays [32]. Abundance o CO2in he a mosphe e du ing ha ime would be a consequence o a ca bon-silica e cycle du ing millions o yea s ha a e changed he Snowball e en s condi ions [33]. The concen a ion o oxygen was low in he oceans du ing he Snow- ball pe iods. The wa e had a high le el o acidi y due o he high concen a ion o CO2 in he a mosphe e [32]. The ice-co e ed condi ions on Ea h we e al e ed because o he mel ing o he ice c us ha ook place due o he inc ease o he empe a u e by olcanoes ac i i y, which educed he p esence o CO2in he a mosphe e and p o- oked he eme gence o liquid wa e [34]. The high olcanic ac i i y igge ed he ex ensi e p esence o hyd o he mal en s du ing he C yogenian age [35–37]. Nowadays, hyd o he mal sys ems can be classi ied as black smoke s and los ci y sys ems. The i s one, a e cha ac e ized by he black smoke ha ises om he chimney-like ocky o ma ions, whe e seawa e is in con ac wi h he magma cham- be s and eme ges wi h an acid pH 2–3, a high con en o dissol ed me als such as Fe (II) and Mn (II), a a ie y o gases o igina ed om olcanic ac i i y like CO2,H 2S, H2, CH4and also wi h high empe a u es up o 405°C. In con as , in he los ci y sys ems, he wa e ha ci cula es ough he en s is no in con ac wi h he magma, ins ead, i is hea ed by con ec ion om he man le and by exo he mic chemical eac ions be ween he luid and he su ounding ocks eaching empe a u es o 200°C [35]. The ock ha in e ac s wi h he luid is domina ed by low-silica i on and magnesium ich mine als, p o oking he me hanogenesis by se pen iniza ion o he hyd ogen and he educ ion o ca bon dioxide in he ocean. In his case, he pH o he luid is basic 9–11, i has dissol ed gases likeH2,CH 4, low-mass o hyd oca bons, and a low dissol ed CO2. Simila i ies could be ound along wi h he ancien oceans on Ea h du ing he Snowball E en s and he cu en condi ions o he ocean on Enceladus. He e we p esen a compa a i e geochemis y analysis o bo h oceans. We also desc ibe a chemical me abolic p ocess based on nume ical simula ions ha could ake place wi hin he global ocean o Enceladus, in o de o in e i he cu en condi ions o ha 3 Is he Ocean o Enceladus in a P imi i e E olu iona y S age? DOI: h p://dx.doi.o g/10.5772/in echopen.104862 ocean could e ol e o c ea e he building chains o li e. Du ing glacia ions ages, he ice-co e ed Ea h allowed o main aining he liquid wa e benea h he ice c us , and subsequen ly ha liquid wa e eme ged o he su ace by he ho spo s o hyd o he - mal en s once he high concen a ion o CO2s a ed changing he condi ions o he a mosphe e [38]. On Enceladus, he e a e hin s ha indica e he p esence o liquid wa e , such as he hyd a ed sodium sal s de ec ed by he Cassini mission. The molec- ula hyd ogen ound also gi es clues abou a hyd o he mal ac i i y benea h he ice shell. We aim o in e a possible e olu iona y s age o he ocean o Enceladus ha could make possible he eme gence o li e. 2. S udy a ea Models o he in e nal s uc u e o Enceladus e eals an ocean on a e age 26– 31 km in dep h below an ice laye be ween 21 and 26 km o hickness [39]. Salini y geochemis y simula ions o he ocean o Enceladus show alues nea ly simila o Ea h, a ound 20 g/kg [40, 41]. The quan i y o wa e apo ejec ed om he plumes is a ound 150–300 kg/s [42]. This ejec ion o pa icles supply he composi ion o he ing E o Sa u n, wi h <10% o he ma e ial ca ching in o i , also sugges ing a liquid o igin [43, 44]. Figu e 1 shows he dis ibu ion o he plumes along he sou h pole o Enceladus. Figu e 1. Digi al ele a ion model (DEM) o he plumes called “Tige s ipes”loca ed in he sou h pole o Enceladus. I was used he images aken by he Cassini Mission. This DEM was de eloped using he so wa e TopoCal 2022 .9.0.811. 4 As onomy and Plane a y Science - F om C yo olcanism o Black Holes and Galac ic E olu ion Benea h he sou h pole he composi ion o he pa icles is mainly sal ich, implying ha hose sal s a e la ge han sal -poo g ains and hey a e expelled wi h lowe escape eloci y. The escape speed o pa icles om he plumes in Enceladus is on a e age 1.85–2.25 km/s, acco ding o he measu es om he dus y plume by he lyby o he Cassini spacec a [45]. Figu es 2 and 3show he longi udinal (y axis) and ans e sal (x axis) heigh p o iles o he plumes o Enceladus om Figu e 1. The longi udinal axis o Figu e 2 p esen s a adius in he cen al plume o 190 m, besides, he ans- e sal axis o Figu e 3 shows a adius o 90 m. The dis ibu ion o he issu es along he plumes seems o be aligned in he y axis. 3. Ma e ials and me hods In his esea ch, we used he da a o he molecula species de ec ed by he INMS ins umen on boa d o he Cassini Mission. The spec al signa u es we e encoded acco ding o Ramí ez-Juidías e al. [46]. Then, he e we e selec ed he common spec al lines p esen in he ocean o Enceladus and in he seawa e o he oceans on Ea h. Based on he spec al lines, i was applied da a mining in o de o ex ac he Figu e 2. Ele a ion p o ile o he DEM om op o bo om. Figu e 3. Ele a ion p o ile o he DEM o he cen e om le o igh . 5 Is he Ocean o Enceladus in a P imi i e E olu iona y S age? DOI: h p://dx.doi.o g/10.5772/in echopen.104862 concen a ion o species de ec ed in he ma e ial ejec ed om he plumes. Table 1 shows some o he species p esen in he ocean o Enceladus and he seawa e o he oceans on Ea h [47] wi h hei concen a ion in g/kg. Each specie was ex apola ed o he geochemical p ocesses associa ed o he ac i i y o CO2and H2O wi hin he ocean [22]. Acco ding o he me hod pa en ed by Ramí ez-Juidías e al. [46], he da a mining p ocess was ca ied ou h ough he applica ion o modi ied gene ic algo i hms, i e a- i ely analyzing a la ge amoun o da a h ough a p ocess simila o gene ic mu a ion, in o de o ex ac he a iables ha a e hen used o ob ain he concen a ions (g/kg) o species in he ocean o Enceladus, using he wa eleng hs be ween 0.35 and 1 μm om he spec al da a aken by he VIMS ins umen . The encoding model de eloped o ob ain hese concen a ions consis s in building a ec o o size equals o he numbe o i e a ions o execu e. The k h-o de o he ec o ep esen s he wo k ha is done in he k h-posi ion. In his case, a popula ion o al e na i e solu ions is se led o a ce ain numbe o ch omosomes, ha ep esen he na u al sequence in which he a iables (spec al signa u es) a e p og ammed. The p ocess o planning and p og amming equi ed o he ex ac ion o he concen a ions o species is usually conduc ed by applying a h ee-le el model called espec i ely S a egic App oach, Tac ical App oach and Ope a ional App oach. This model can be eplica ed using machine lea ning. Species Enceladus concen a ion (g/kg) Ea h concen a ion (g/kg) BOHðÞ 40,008 BOHðÞ 30,019 B 0,067 Ca2þ0,412 Cl7076 19,353 CO322867 0,016 F0,013 HCO30,031 0,107 Kþ0,399 Mg2þ1284 Naþ7343 10,784 NaCl 0,024 NaCO31788 NaHCO30,015 NaOH 0,008 OH0,038 SO420,1–0,01 2713 S 2þ0,008 Table 1. Concen a ion o species in he ocean o Enceladus and in he seawa e o he oceans on ea h. 6 As onomy and Plane a y Science - F om C yo olcanism o Black Holes and Galac ic E olu ion 4. Resul s and discussion Sodium ion and Chlo ine a e he mos abundan species in he ocean o Enceladus. Figu e 4 shows he concen a ion o bo h species in mol pe kg o H2O. The quan i y o mol is calcula ed in unc ion o he CO2ac i i y. Naþis p esen in concen a ions a ound 0.800 mol/kg while he concen a ions o Cla e cons an , a ound 0.400 mol/ kg. Figu e 5 also shows ha he e a e p esen some ca bona es CO32, bica bona e HCO3and sul a e SO42.CO 32has a concen a ion on a e age 0.075–0.030 mol/kg which ends o dec ease wi h he ac i i y o CO2. HCO3p esen s an inc emen om 0.020 o 0.070 mol/kg and SO42has a concen a ion be ween 0.01 and 0.1 mol/kg. The concen a ions o salini y and chlo ini y a e ela i ely cons an in he cu en e es ial oceans. The a e age concen a ion om he seawa e wi h a pH o 8.1 and empe a u e o 25°C a e de ailed in Table 1. Geological in o ma ion ex ac ed om sedimen a y laye s e eals ha deep oceans we e in a educed s a e ill he end o he Paleop o e ozoic e a. I on and calcium sul a e p obably played as educed agen s wi h he oxygen con e ing FeO in o Fe2O3, and p ecipi a ing CaSO4. Du ing he Figu e 4. Sodium ion and chlo ine p esen in he ocean o Enceladus. The concen a ion o species a e calcula ed in unc ion o he CO2ac i i y. Figu e 5. Some ca bona es, bica bona es, and sul a e p esen in he ocean o Enceladus. The concen a ion o species a e calcula ed in unc ion o he CO2ac i i y. 7 Is he Ocean o Enceladus in a P imi i e E olu iona y S age? DOI: h p://dx.doi.o g/10.5772/in echopen.104862 C yogenian e a he concen a ion o sulpha e could ise o le els simila o he ecen ones, a ound 23 mol/kg o H2O [47]. Table 2 shows ew key species p esen in he ocean o Enceladus and in he seawa e o he oceans on Ea h. Sodium ion and Chlo ine a e he mos abundan species in bo h oceans. The oceans on Ea h a e sal ie wi h a pH o 8.1 on a e age, while he ocean o Enceladus is mo e basic, a ound pH 12.2. The ocean o Enceladus has mo e dissol ed ino ganic ca bon han he ocean on Ea h. On Enceladus, he p edominan ca bona e is CO32while on Ea h is he bica bona eHCO3. The abun- dance o CO32in he ocean o Enceladus could be due o he se pen iniza ion o he molecula hyd ogen. The concen a ion o sul u in he ocean o Enceladus is a iable compa ed o he one p esen on Ea h. Two scena ios can be conside ed o calcula e he amoun o sulpha e ha could be oxidized on he ocean o Enceladus. The lowe concen a ion o SO42, 0.01 g/kg, displayed in Table 2, akes place only in aqueous educ an s en i onmen s whe e HS eac s wi h he oxidan s, while in he la ge concen a ion 0.1 g/kg, some mine als a e conside ed as a sou ce o educ an s. The concen a ion o sulpha e in he ocean o Enceladus is below o he cu en amoun o sulpha e ound on he oceans on Ea h bu , his concen a ion could ha e been smalle du ing he Snowball e en s, being close o he cu en quan i ies on he ocean o Enceladus. The p edominan concen a ion o ino ganic ca bona e species ound in he ocean o Enceladus, se he ocean as no compa ible wi h li e excep o he me hane de ec ed ha can be a p oduc o he me hanogenesis o he ca bon dioxide and he hyd ogen. Would i be possible ha he species de ec ed in he ocean o Enceladus e ol e o c ea e he chains o li e? how we e he chemical condi ions o he p imi i e e es ial oceans be o e ising li e? In o de o igu e ou which simila i ies could be ound be ween he e es ial oceans and he ocean o Enceladus, i is necessa y o unde s and he e olu ion o he ancien aqueous geochemis y o he oceans in he p imi i e Ea h. Du ing he i s s age o o ma ion o Ea h, i was bomba ded by hyd ous as e - oids mainly ype Cl chond i es b inging wa e , o ganic molecules, and chond i ic mine als. Tec onic ac i i y acili a ed o di e si y he mine alogy along he c us , inc easing he ma ic con en o he op laye s h ough he e up ion o ho basal ic la as. Chond i ic ma e ial has been also de ec ed in he plumes o Enceladus [14, 21], ha is why, i could be possible o in e ha his ma e ial can be se led in he sea loo o i s ocean [48]. O ganisms canno de ise chemical p ocesses by hemsel es, hey mus copy na u- al eac ions, adap hem, and op imize hem h ough ime. Phospho yla ion is he addi ion o a phospha e g oup in o a p o ein, being he main mechanism o Species Enceladus concen a ion (g/kg) Ea h concen a ion (g/kg) Cl7076 19,353 CO322867 0,016 HCO30,031 0,107 Naþ7343 10,784 SO420,1 - 0,01 2713 Table 2. Concen a ion o species p esen in he seawa e o he oceans on ea h and in he ocean o Enceladus. 8 As onomy and Plane a y Science - F om C yo olcanism o Black Holes and Galac ic E olu ion [44] Juhász A, Ho ányi M. Seasonal a ia ions in Sa u n's E- ing. Geophysical Resea ch Le e s. 2004;31: 19703. DOI: 10.1029/2004GL020999 [45] Sou hwo h B, Kemp S, Schmid J. Modeling Eu opa's dus plumes. Geophysical Resea ch Le e s. 2015;42: 541-548. DOI: 10.1002/2015GL066502 [46] Ramí ez-Juidías E, Pozo-Mo ales L, Galán-O iz L. P ocedu e o ob aining a emo e sensed image om a pho og aph. Pa en n° ES2537783B2 (2015-09-29 publica ion o he pa en concession). In e na ional Pa en n° WO201419897 4A1. Uni e sidad de Se illa. 2015 [47] Mille o F, B uland K, Lohan M, Nigh ingale P, Liss P, de la Rocha C, e al. The oceans and ma ine geochemis y. In: Elde ield H, edi o . T ea ise on Geochemis y. Vol. 6. Ox o d, UK: Else ie ; 2003 [48] San osh M, A ai T, Ma uyama S. Hadean ea h and p imo dial con inen s: The c adle o p ebio ic li e. Geoscience F on ie s. 2017;8:309-327. DOI: 10.1016/ j.gs .2016.07.005 [49] Giba d C, Go ell I, Jiménez E, Kee T, Pasek M, K ishnamu hy R. Geochemical sou ces and a ailabili y o Amidophospha es on he ea ly ea h. Angewand e Chemie. 2019;131: 8235-8239. DOI: 10.1002/ ange.201903808 [50] Va gas M, Kashe i K, Blun -ha is E, Lo ley D. Mic obiological e idence o Fe(III) educ ion on ea ly ea h. Na u e. 1998;395:65-67. DOI: 10.1038/25720 [51] Deame D, Dame B. Can li e begin on Enceladus? A pe spec i e om hyd o he mal chemis y. As obiology. 2017;17:834-839. DOI: 10.1089/ as .2016.1610 [52] McKay C, Anba A, Po co C, Tsou P. Follow he plume: The habi abili y o Enceladus. As obiology. 2014;14: 352-355. DOI: 10.1089/as .2014.1158 [53] Po co C, Dones L, Mi chell C. Could i Be snowing mic obes on Enceladus? Assessing condi ions in i s plume and implica ions o u u e missions. As obiology. 2017;17:876-901. DOI: 10.1089/as .2017.1665 [54] Taubne R, Pappen ei e P, Zwicke J, Sm zka D, P uckne C, Kola P, e al. Biological me hane p oduc ion unde pu a i e Enceladus- like condi ions. Na u e. 2018;9:1-11. DOI: 10.1038/s41467-018-02876-y 15 Is he Ocean o Enceladus in a P imi i e E olu iona y S age? DOI: h p://dx.doi.o g/10.5772/in echopen.104862