Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , 1

Open Access , J. Chen 1 Discussions , Chemistry and Physics Atmospheric , R. Mason 1 1 ect on the Eurotiomycetes ff , M. J. Wheeler 1,* 5014 5013 and Agaricomycetes , and A. K. Bertram 3 , R. Iannone 2 , U. Pöschl 1 , S. M. Burrows 1 This discussion paper is/has beenand under Physics review (ACP). for Please the refer journal to Atmospheric the Chemistry corresponding final paper in ACP if available. Department of Chemistry, University of British Columbia, 2036 MainAtmospheric Mall, Sciences Vancouver, British and Global Change Division, Pacific Northwest National Laboratory, Max Planck Institute for Chemistry, Hahn-Meitner-Weg 1, 55128 Mainz, Germany now at: SNC-Lavalin Environment, 8648 Commerce Court, Burnaby, British Columbia, Received: 20 December 2013 – Accepted: 22Correspondence January to: 2014 A. – K. Published: Bertramand 21 ([email protected]) S. February M. 2014 Burrows ([email protected]) Published by Copernicus Publications on behalf of the European Geosciences Union. Atmos. Chem. Phys. Discuss., 14, 5013–5059,www.atmos-chem-phys-discuss.net/14/5013/2014/ 2014 doi:10.5194/acpd-14-5013-2014 © Author(s) 2014. CC Attribution 3.0 License. atmospheric transport of fungal spores from the classes Ustilaginomycetes, E. A. Polishchuk 1 Columbia, V6T 1Z1,2 Canada P.O. Box 999, Richland,3 Washington, 99352, USA * V5A 4N6, Canada Ice nucleation and its e D. I. Haga Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Eu- cient ffi C, and ◦ , and 31 have been − 1 − . We show that erent species of , which together ff C and ◦ man et al., 2013; Tobo ff 25.5 − Ustilaginomycetes , Ascomycota Eurotiomycetes ' and have been reported in the continental 1 − ) there is a wide range of freezing proper- C, 0.01 between ◦ Agaricomycetes ect the frequency and properties of ice and 5016 5015 ff 29 − Agaricomycetes Basidiomycota are found on all types of decaying material and in- > C. On average, the order of ice nucleating ability for C and include many types of mushroom species and are cos- ◦ n, 2004). ◦ are agricultural pathogens and have caused widespread Basidiomycota ffi C, all of the fungal spores studied here are less e ◦ 36 19 − − 20 and − C and Eurotiomycetes ◦ Ustilaginomycetes 26 Agaricomycetes − . Ascomycota Ustilaginomycetes Ice nucleation on fungal spores may be important because ice nucleation on these et al., 2013; Hader etbecause al., 2013). ice Ice nucleation nucleation on on fungal fungal spores spores, may followed also by be important precipitation, may be an impor- important than ice nucleation on mineralbut dust ice (Hoose nucleation et on al., fungal 2010; sporesice Sesartic could and et al., still mixed-phase 2013), clouds influence the regionallyPrenni, frequency and 2010; and seasonally properties Möhler (Delort et of et al., al.,et 2007; 2010; al., DeMott Morris 1982; and et Phillips al., etAs 2004, examples, 2011; al., numerous Pöschl 2009; field et studies Creamean have al., et illustratedconditions, 2010; that al., a Sands at 2013; certain fraction times, Spracklen of locations and(Pratt the and Heald, et total 2013). al., ice 2009; nuclei Prenni population et may al., contain 2009; biological Garcia material et al., 2012; Hu 1955; Proctor and Parker,Imshenetsky 1938) et al., and 1978; even Gri into the stratosphere (Smithspores could et influence al., the 2010; frequencythe and atmosphere. On properties a of global ice annual and scale, ice mixed-phase nucleation clouds on in fungal spores may be less Fungal spores, which are thedant reproductive structures in in the the atmosphere lifecycleerage (Després of fungal fungi, et spore are al., concentrations abun- 2012; ofboundary Madelin, 1 layer 1994; to (Elbert 10 Bauer et L et al.,previously al., 2007) observed and 2008). (Oliveira peak et Av- al., concentrations 2005; oftero Ho 20 et et to al., al., 35 2005; L 2010; Goncalves etcan Khattab al., be and 2010; transported Quin- Levetin, large 2008; distances(Hirst Nayar and et and transported al., to Jothish, 1967; high 2013). altitudes Gregory, These 1978;1989; in Bowers the spores Fulton, et troposphere al., 1966; 2009; Kelly Amato et and al., Pady, 2007; 1953; Ebner et Pady al., and Kelly, 1953; Pady and Kapica, decrease annual mean mixing ratios in the upper troposphere. 1 Introduction trations of fungal spores in near-surface air over the oceans and polar regions and peratures of spores frommake the up phyla 98 %both of phyla ( known fungalties, species and found also on thatthe the Earth. average variation freezing The within properties data aport of phylum show model, the is we that phyla. greater investigated within Using than whetherprecipitation, a ice the can global nucleation influence variation on chemistry–climate the between the atmospheric trans- spores studied transport in the spores, and atmosphere. followed global Simulations by distributions showof that of these inclusion these of fungal ice spores nucleation in scavenging mixed-phase clouds can decrease the annual mean concen- 0.1 between these spores is at temperatures below ice nuclei compared tonew Asian freezing mineral results dust together on with a data per in surface the area literature basis. We to used compare our the freezing tem- mopolitan. damage to crops. clude important human allergens. Weto focused on be these abundant classes in sinceice they the nucleation are atmosphere ability thought of and these because classesinvestigated of there were spores is found in very the to literature.than little cause All homogeneous information of freezing. freezing the on The of fungal cumulative the spores numberat water of temperatures droplets ice between nuclei at per temperatures spore warmer was 0.001 Ice nucleation on fungalmixed-phase clouds. spores We studied may the a icefungal nucleation spores properties chosen of from three 12 classes: di rotiomycetes Abstract 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | that (Mag- Agari- 1 − Aspergillus Agaricus bis- to 1 L have also been and 3 − Eurotiomycetes spore concentrations . 1 − genera. These spores are . The species and classes of Eurotiomycetes Agaricus bisporus (Nayar and Jothish, 2013; Magyar 1 Agaricomycetes − Aspergillus and 5018 5017 erent classes (Hibbett et al., 2007). So far, the ff Penicillium erent species of fungal spores chosen from three classes: ff Exobasidiomycetes, Eurotiomycetes, Pucciniomycetes, Sor- , spores have frequently been identified from surface air samples are agricultural pathogens that belong to the group of smut fungi, is a class of fungi that includes many types of mushroom species are found on all types of decaying material and are one of the most Ustilaginomycetes, and Eurotiomycetes , (Haga et al., 2013; Morris et al., 2013b; Iannone et al., 2011; Jayaweera . The few studies that have quantified Dothideomycetes , the common or button mushroom. Spores from this class have been found to ). Our studies are complimentary to the studies by Pummer et al. (2013) since Eurotiomycetes Ustilaginomycetes Agaricomycetes Fungi can be classified into 35 di et al., 2010; Mallo et al., 2011; Shelton et al., 2002; Fröhlich-Nowoisky et al., 2009), causing widespread damage to cropsunderstanding and cereals the (Webster transport and of Weber, 2007). thesenucleation) Hence, spores is in of the interest atmosphere from (which an may economic involve perspective. ice ubiquitous types of fungi (Websteridentified and as Weber, 2007). important humanreferences therein). allergens The (see, specific forwere types example: studied of Horner here are spores etfrequently from from al. the present the (1995) in class and surface air (Pyrri and Kapsanaki-Gotsi, 2012, 2007; Goncalves et al., 2009;et Khattab al., and 2006; Hasnain Levetin, etet 2008; al., al., 2005; Pyrri Wu 2000; et and Calderon al.,class et Kapsanaki-Gotsi, have 2004; been al., Troutt 2007; identified and 1995; at Levetin, Morales high 2001; Hirst,Ustilagomycycetes altitudes Sabariego 1953; in the Gregory, troposphere 1952), by Pady and and Kelly spores (1954). from this comycetes to the genus level have reportedyar et surface concentrations al., of 2009; roughly Morales 10 et al., 2006; Li, 2005). (Mallo et al., 2011; Pyrri andto Kapsanaki-Gotsi, a 2007) third and of have the been total2006; shown fungal Hasnain to et spores make al., in up 2005; some Mitakakis regionshave and (Herrero been Guest, et 2001). al., measured Boundary 2006; to layer Morales concentrations et be al., roughly 0.05 to 6 L (Fröhlich-Nowoisky et al., 2012;et Mallo al., et 2009a, b; al.,2006; Magyar Herrero 2011; et et Yamamoto et al., al.,et 2006; al., 2009; al., Morales 2012; Fröhlich-Nowoisky 2009; et et Amato Oliveira fied al., et al., from 2006), al., 2009; continental and 2007). air Zoppas by For also et Fröhlich-Nowoisky example, at et al., over high al. half (2012) altitudes of were (Bowers the from fungal the class species identi- be important components of the air-borne fungal spore population near the surface porus are thought to be abundanta in the few atmosphere. of Recently, Pummer theniger et same al. species (2013) studied aswe were measured studied the here cumulative ( numberwell of as the IN ice per nucleation spore activePummer as surface et site al. a (2013) density function reported as of the ausing temperature temperature, function an at of as average which temperature, spore 50 while % mass of concentration all of of about the 20 droplets mgmL froze and are cosmopolitan (Webster and Weber, 2007). This class includes and Flanagan, 1982; Pouleur etamount of al., data 1992; on Pummer the et icecleation nucleation al., properties properties 2013). of of To 12 fungal add spores, di Agaricomycetes to we studied the the limited ice nu- fungal spores investigated are listed in Table 1. We focused on these classes since they Pfender et al., 2006;but Skelsey ice et nucleation al., followed by 2008;of precipitation Wang the was et spores not al., from included 2010; in thebe these atmosphere. Wilkinson important, models Even very et as though little al., a ice is 2012), sink known nucleation about on this fungal topic. sporesice may nucleation propertiesgated: of sporesdariomycetes from the following classes have been investi- The main mechanism for thester spread and of Weber, fungi 2007), is and by accurateneeded. wind-dissemination models In to of the predict spores past the (Web- researchersious spread have of models predicted fungal (Andrade the spores spread et are ofIsard al., fungal 2009; et spores Aylor, al., using 1986, var- 2005; 2003; Kim Fröhlich-Nowoisky et and al., Beresford, 2012; 2008; Magarey et al., 2007; Pan et al., 2006; tant removal mechanism of these spores from the atmosphere (Morris et al., 2013a). 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | 1 − Pen- Agari- Agari- Basid- ) were (collec- ) spores. in remote ) were ac- cient ice nu- ffi Agaricomycetes spores have also make up approxi- spores being more Ascomycota. Basidiomycota spores as high as 10 L Pucciniomycetes Ustilago avenae spores (Amato et al., 2007; to , Trichaptum abietinum Penicillium spores were prepared analo- sp. (collection #58), and , Aspergillus brasiliensis Ascomycota spores are more e spores were isolated using tech- , we also investigated whether ice Basidiomycota Penicillium and (Nayar and Jothish, 2013; Fernández Aspergillus Ascomycota 1 Penicillium Ustilago nigra − , Lepista nuda Aspergillus , 5020 5019 Basidiomycota Ustilaginomycetes Eurotiomycetes Basidiomycota n, 2004; Jayaweera and Flanagan, 1982; Imshenet- spores, resulting in ffi , and Ustilago nuda (collection #161), Boletus zelleri , (collection #54). species studied here were obtained from the UBC Depart- spores ( , the common button mushroom, was purchased from a local grocery Ascomycota Ustilaginomycetes spores, sections of the fruiting body were placed on a wire mesh in a sealed , Aspergillus niger are better ice nuclei than fungal spores from the phylum Eurotiomycetes Amanita muscaria ciently removed from the atmosphere by precipitation from ice and mixed-phase Ustilaginomycetes The Fungal spores from the phyla In addition to studying the ice nucleation properties of spores from the classes ffi cillium brevicompactum niques similar to those described by Jones et al. (1992) and Allan and Prosser (1983). gously to the methodBriefly, used a by sealed Haga glassa et sonicator vessel al. bath. containing The (2013) combination spores forgenerated of a rust the and spore fan ( cloud a and within the smallglass the vibrations slides vessel from fan that and the were were resulted sonicator suspended in bath immersed on spores in a being wire deposited mesh onto ment within the of vessel. Chemistry Bioservicestion Laboratory #56), collection: mesh, exposed to the parttions. Over of time, the spores fungus were that naturally releases released spores and under deposited onto humidifiedquired the from condi- the glass Cereal slides. Research Centre,Manitoba, Agriculture Canada. and Agri-Food Slides Canada, containing Winnipeg, 2.1 Spore samples and slide preparation Agaricus bisporus store in Vancouver, Britishfungi Columbia (B.C.), ( Canada. Theharvested remaining from the PacificColumbia Spirit (UBC) Regional Park campuscomycetes surrounding in the Vancouver, Universityand of B.C. humidified British To chamber. Hydrophobic prepare glass slides slides were containing placed underneath the wire iomycota 2 Methods with freezing data from the literature to assess if fungal spores from the phylum the marine boundary layer, polar regions, and the upper troposphere). mately 98 % offield known measurements observed fungal higher species ratiosmarine of found regions compared on to Earth continentalpossible regions (James explanation (Fröhlich-Nowoisky for et et this al., al., wasclei 2012). that 2006). One compared Recent to e clouds (Fröhlich-Nowoisky et al., 2012). We also use our new freezing data, together nucleation on these spores,removal followed by of precipitation, these canclimate influence spores transport the in model transport and the and byincluded atmosphere. comparing in simulations We the with did and model.studied without this may These ice be using simulations nucleation important a suggest for the that global transport ice chemistry- of these nucleation spores on to the remote regions spores (such as been identified at2010; high Amato altitudes et insky al., the et 2007; al., troposphere 1978; Gri Fulton, andPady, 1966; 1953; Pady Proctor stratosphere and and Kapica, (Smith Parker, 1938), 1955; asImshenetsky et Pady have and et al., Kelly, al., 1954; Kelly 1978; and 1938). Fulton, 1966; Pady and Kapica, 1955; Proctorcomycetes and Parker, surface concentrations of roughlyet 0.1 al., to 2012; 5 Crawford L 2010; et Khattab al., and 2009; Levetin, 2008;riods Pyrri Wu of and et high Kapsanaki-Gotsi, spore al., 2012; productivity, 2004;have concentrations Quintero Li been of and observed et Kendrick, (Pyrri al., 1995). and During Kapsanaki-Gotsi, pe- 2012). and they often represent a large majority of identified spores (up to 35 %) with typical 5 5 20 25 15 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , , , Pencil- U. ave- Penicillium A. bisporus A. bisporus glass beads A. muscaria , and ) spores were A. brasiliensis 3 objective) cou- × and Bertram, 2010; spores ( U. nigra , ff U. nigra or 50 spores ( , × U. nuda , C. Finally, the temperature was ◦ U. avenae Aspergillus 40 , xed onto aluminum SEM stubs, sput- − Agaricomycetes C and the temperature of the flow cell ◦ ffi A. niger , ) and spores on hydrophobic surfaces (Iannone U. nuda (aq)), and then a portion of the culture was ( spores were imaged by field emission SEM 4 5022 5021 down to 1 − T. abietinum A. brasiliensis , C min , ◦ Cladosporium samples for imaging, cultures of each fungal species were xed onto aluminum SEM stubs using double-sided glue tabs, ffi L. nuda Ustilaginomycetes , P. brevicompactum T. abietinum , ) were a B. zelleri , sp., and L. nuda , P. brevicompactum spores were aerosolized from fungal cultures and deposited on a hydrophobic C. Next, the flow cell was isolated from the humidified flow and the temperature A. bisporus were acquired using variable pressure SEM (Hitachi S2600 VP-SEM). ◦ Scanning electron microscopy (SEM) images, acquired at the UBC Bioimaging Facil- chemistry-climate model,EMAC ECHAM/MESSy model is Atmospheric a numerical Chemistry chemistry (EMAC). and climate The simulation system that includes first fixed using osmiumexcised vapor (4 and % mounted OsO ontoadhesive), an and aluminum finally the SEM samples stub wereexperiments, allowed using images to were Aquadag air captured dry (a using prior graphite secondarytween to based electrons, 5 imaging. a For and working the 15 distance SEM mm, be- and an acceleration voltage2.4 between 3 and 25 Global kV. chemistry-climate transport model, EMAC We simulated the transport and removal of atmospheric aerosol particles in a global sputter coated with 12ment. nm Similarly, gold loose (Au) palladiumdeposited directly (Pd) onto and double-sided glue mounted tabs,ter a in coated the with SEM 8 nm instru-sp. and Au and mounted in the SEM instrument. To prepare ity, were used to determineB. the zelleri surface features of thenae spores. Images for Penicillium (Hitachi S4700 FESEM). Glass slidesA. containing muscaria and Using microscope images, wespore calculated and the the average projected sporemation 2-D dimensions and surface (length area the and per width) 2-Dnumber for of surface each spores area per species. of drop. From spores this infor- per drop (Sect. 2.2), we determined the 2.3 Spore properties and number of spores per drop surface area of the fungal spores contained within each droplet were determined. Wheeler and Bertram, 2012),glass and cover slide will is only usedvapor be to is described support condensed briefly the onby fungal here. the adjusting spores A spores the inside dew hydrophobic to point the grow ofto flow the droplets 0 cell. carrier approximately First, gas 30–150 water to µmwas 2 in lowered size at araised rate to of 5–10 room temperatureon to the evaporate the glass droplets, slide.experiments. leaving For Digital only each video the droplet, and fungal the temperature spores freezing were temperature recorded and the during 2-dimensional the (2-D) freezing The instrument used toconsisted study of the an optical immersion microscope freezingpled (Zeiss with properties Axiolab a A of flow with the cell aor that fungal 10 a had spores similar temperature apparatus and has relativeet been humidity used al., control. in This 2011; many previous apparatus Dymarska ice nucleation et studies al., (Iannone 2006; Koop et al., 1998; Cherno settled to the bottomonto of hydrophobic the glass sample vial. slideslium The using resulting a spore Meinhard suspension nebulizerglass was (model slide sprayed using TR-30-A1). a custom-builtviously flow used cell recently for developed depositing et in al., our 2011). laboratory and pre- 2.2 Freezing experiments In short, the(0.5 mm fungi in were diameter) (BioSpec grown Productsculture, Inc.) in which were rolled petri resulted over in theto dishes surface the the of and the beads. spores approximately fungal The beinging 10 beads dislodged ultrapure and from water. spores the This were culture resulted then and transferred in attached to a a suspension sample of vial spores, contain- while the glass beads 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | is er- ff 25 and nuc, liq − R 210 km at cients used × ffi are the fraction value of zero at C), and in the sec- ◦ nuc, ice nuc, ice 35 R is zero for mixed phase R − and C) we assume that nuc, ice ◦ R 0 nuc, liq > R 1. For mixed-phase clouds (i.e. clouds = , where ), due to nucleation scavenging and subse- R i C) we carried out simulations with two di 5024 5023 er between IN-Active and IN-Inactive, the sen- ◦ X nuc, liq ff erent simulations were carried out to test if ice R 35 ff − C for the onset of mixed-phase ice nucleation scaveng- ◦ C, but equal to unity for temperatures between ◦ C and ◦ 25 25 − C may also di ◦ − is 0.05 for both simulations mentioned above (IN-Inactive and in latitude and longitude, or approximately 140km ◦ > 35 . These two di cient cloud condensation nuclei (CCN). Here, we are using the − ffi nuc, ice R nuc, ice R C, which is the homogeneous freezing temperature of water droplets) ◦ 35 erent simulations is included in Table 2. ff ≤ − C. A temperature of ◦ The in-cloud local rate of change in the mixing ratio of particles, i.e. the number of Removal processes of particles simulated in the model included sedimentation, dry Simulations were performed for five years and an additional year of spin-up time, with 35 particles per number of air molecules ( atures we assume that IN-Active). This valuetemperatures is (Henning consistent et with al.,temperatures 2004; measurements below Verheggen of et cloud al.,sitivity 2007). scavenging of Although modeled at scavenging aerosol low at transportlow to (Burrows scavenging et at al., these 2013).in temperatures the A is di summary extremely of the nucleation scavenging coe ond simulation (referred to asclouds IN-Active) at we temperature assumed − ing was chosen becauseoccurs for it the represents spores the studied approximate here (see temperature Fig. where 3). freezing For ice clouds (i.e. clouds at temper- order for the sporesincorporated to in liquid act cloud as dropletswhich immersion-freezing during is the nuclei, ensured droplet they nucleation byat would and the temperatures growth first assumption process, between have 0 toent be values of nucleation can impact theIn long-distance the transport first of simulation theall (referred particles temperatures to relevant in as for the IN-Inactive) mixed-phase atmosphere. we clouds used (below an 0 to of particles embedded inFor warm cloud liquid droplets clouds orunity, (i.e. as ice clouds done particles at previously temperatures (Heald withinthe and a particles Spracklen, cloud, 2009). are respectively. Aimmersion e value freezing of results unity to assumes assess that implications for fungal spore global transport. In prescribed with a scavenging parameter, sedimentation, dry deposition, and impaction scavenging. Nucleation scavenging was Earth’s surface. Atmospheric transportterized is simulated convective mass (including advection flux),the and and parame- surface. tracers The are EMAC removedcapable model, by of used realistic dry in simulations and similar ofAfrican wet configurations, aerosol dust deposition has transport to to and been Europe deposition shownthe (Gläser for Chernobyl to the et accident be transport (Lelieveld al., et of 2012) al., and 2012). of radioactivedeposition, aerosol impaction particles scavenging, from andand nucleation scavenging ice by clouds. liquid, We mixed-phase, use the methods outlined by Burrows et al. (2009) to describe the model setup described inthe detail scavenging by scheme Burrows (Tost et etcrystal-borne al. species al., (2009), due and 2010), to with including the modificationstitioning the sedimentation to of slow of tracers crystals, downward associated and transportBriefly, with the aerosols of evaporation release of are and droplets treated repar- and as melting monodisperse of passive ice aerosol crystals. tracers, emitted at the EMAC (ECHAM5 version 5.3.01,with MESSy a spherical version truncation 1.9)proximately of in 1.9 T63 by the (corresponding 1.9 T63 to L31mid-latitudes) a resolution, with quadratic i.e. 31 Gaussian vertical gridlution of hybrid provides ap- a pressure reasonable levelstracer balance up transport between to in accuracy the 10 hPa. ECHAM5 andto GCM This computational further has expense: model increases been in reso- shown model to resolution be (Aghedo comparatively et insensitive al., 2010). teraction with oceans, land andet human al., influences 2006; (Jöckel Tost et etSubmodel al., al., 2006, System 2006; 2005; Lawrence (MESSy) Kerkweg andmospheric to Rasch, model link 2005). is It the multi-institutional usesmodel 5th computer the generation (ECHAM5 Modular codes. European GCM) Earth Centre The Hamburg (Roeckner core general et circulation at- al., 2003). For the present study, we applied sub-models describing tropospheric and middle atmosphere processes and their in- 5 5 25 15 20 10 25 20 10 15 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | C T. A. ◦ , and Usti- 19 − Ustilagi- nuc, ice R U. avenae , erent fungal species ff (1981), Mathre (1982), U. nigra , ff ) or as partial spheroids ( U. nuda is the grid-box mean cloud frac- , ) was determined from the number cl f values for the di , (1) ! ciencies of cloud ice and cloud liquid water, A. bisporus 5026 5025 ffi rain F P. brevicompactum erent fungal species studied, conclusions about · spores are shown in blue (open symbols), ff INperSpore(T) lwc sp., nuc, liq R INperSpore(T) + ’s), are previous results from Iannone et al. (2011) for ho- x spores are shown in blue (open blue symbols), snow F Penicillium · spores are in red (open red symbols with vertical line). From , are the fluxes of snow and rain, respectively; iwc and lwc are the Agaricomycetes iwc rain spores are in red (open symbols with vertical line). nuc, ice spores are shown in green (open symbols with horizontal line), and F R ) or from a loss in turgidity of the fungal spores in the vacuum of the SEM A. niger spores are shown in green (open green symbols with horizontal line),

, and 2.3 is the length of the model time step; cl ). These features are likely due to the sample processing prior to imaging f Agaricomycetes t i are the nucleation scavenging e ∆ X snow INperSpore(T) F = Eurotiomycetes i t Shown in Fig. 3 are the The median number of spores per droplet is given for each spore species in Table 1. For nucleation scavenging the same scavenging parameterizations were applied to X nuc, liq ∆ ∆ (see Sect. quent precipitation is: nomycetes and Fig. 3, the cumulative number of ice nuclei per spore was 0.001 between spore as a functionof of temperature spores ( per(Fig. drop 2). and For details, the see fraction Haga et of al. droplets (2013). studied. frozen as a function of temperature shows that the fungal spores studiedtures here compared initiate to the freezing homogeneous at results warmerof of freezing spores Iannone tempera- per et al. droplet (2011). variesthe Since for the relative the number freezing di properties of these spores should not3.3 be formulated from Cumulative Fig. 2. number of ice nuclei per sporeUsing as the a method function presented by of Vali temperature, (1971), the cumulative number of ice nuclei per laginomycetes Eurotiomycetes Also included in Fig. 2mogeneous (as nucleation using the same experimental method as used here. Figure 2 instrument. 3.2 Fraction of droplets frozen vs.Shown temperature in Fig.cluding 2 all is freezing the eventsdroplets that fraction containing were of observed frozen during droplets these as experiments. Results a for function of temperature, in- (1987). SEM imagesthe of images, the the spores fungal appearbrasiliensis spores deflated studied ( areabietinum shown in Fig. 1. In many of Listed in Table 1 are theoptical average dimensions microscopy. Values of reported the in spores studied TableBockus 1 here, et are determined al. consistent by with (2010), theMitchell Gilbertson literature and and values: Ryvarden Walter (1987), (1999), Linco Pasanen et al. (1991), Raper et al. (1949), and Vanky for 3 µm and 8 µmequivalent diameters) diameter of spherical the spores particles, studiedspores which here (see overlaps were Table 1). with emitted In the the globally simulations sizessurfaces, and fungal excluding (volume land continuously ice. at a single constant rate from all land 3 Results and discussion 3.1 Spore properties ice water content andR liquid water content ofrespectively the (and cloud, as respectively; described and in Table 2). both stratiform clouds and to parameterized convective clouds. Simulations were run where tion; 5 5 20 25 15 10 20 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , C P. A. C. ◦ ◦ and C for ◦ 36 spores − U. nigra 27 sp. and − Ustilagino- A. brasilien- ) were grown C and INperSpore(T) P. frequentens culture after 13 ◦ Agaricomycetes and ected by surface ff 26 spores. values for , there was no indi- , the − , and Agaricomycetes Penicillium , Eurotiomycetes U. nigra C for the ◦ T. abietinum 29 A. muscaria P. notatum − values can vary drastically (e.g. P. brevicompactum , results from the current study with A. brasiliensis INperSpore(T) s , values from Niemand et al. (2012) Eurotiomycetes s n sp., , and n C and 0.1 between values for the dust only at temperatures Ustilaginomycetes ◦ . To illustrate this point, as an example, for the Asian mineral dust was calculated value of 0.01 varies by approximately 20 31 P. digitatum cultures indicated surface contamination on L. nuda C for the INperSpore Penicillium − ◦ , 5028 5027 results). Penicillium , 28.5 − values for Asian mineral dusts over the entire tem- C and ◦ U. nuda B. zelleri , INperSpore INperParticle(T) Eurotiomycetes A. niger 25.5 , INperParticle(T) ' Penicillium − C. ◦ U. nigra spores, 0.01 at approximately 25 , − INperParticle(T) are poor IN compared to spores studied here are poorer IN than those studied by Jayaweera spectra presented in the current study should be taken as upper limits. A. brasiliensis U. avenae Agaricomycetes C, 0.01 between ◦ > ) were qualitatively tested for surface contamination by bacteria and/or fungi 29 − Ustilaginomycetes INperSpore cient IN on a per particle basis because they have more surface area available for , all of the spores studied here are poorer IN than the Asian mineral dust studied by Mineral dust particles are considered to be important atmospheric ice nuclei. For It is important to consider the possibility that the fungal spores studied here may con- Jayaweera and Flanagan (1982) and Pummer et al. (2013) are the only other studies From Fig. 3, when comparing the temperatures at which the cumulative number of ffi Since heterogeneous ice nucleation ise a surface process, larger particles may be more sis Niemand et al. (2012)are on similar a to per the particleperature range basis. studied The by Niemandvalues et are al. comparable (2012); to for the below approximately 3.4 Ice nucleation active surface site density, comparison in Fig. 3ing (half-filled results diamonds) by we have NiemandACI04_16 included et and recent ACI04_19). al. immersion (2012) freez- from the for ice submicron nucleation active Asian surfaceand site mineral density, as dust described by (experiments Haga et al. (2013). With the exception of Specifically, the spores after 4 days, anddays. colonies For were the observed remaining in spores, the cation of surface contamination. Fromice the nucleation properties discussion of above, the it sporescontamination seems studied by unlikely here fungi that are or appreciably the bacteria. a However,the we cannot rule out this possibility and, so, bisporus using the same methodto from be Haga et tested al.support were (2013) contaminant cultured for growth in rust andindicated and media the low bunt cultures (nutrient levels spores: were of agar surface the monitored contamination and for spores for two four tryptic of weeks. soy the The fungal agar) test spores studied that here. would culture stage of these experiments. The remaining spores (with the exception of studied here ( in culture, any bacterial contamination of the spores would have been apparent in the the freezing temperature at an in Fig. 3 and looking at the tain bacterial or fungal contaminantsthese on their contaminants surface, would and to changeris discuss the the et likelihood al., ice that 2013b). nucleatingany From properties bacterial the of SEM or images the fungalBioimaging presented spores Facility, surface personal in (Mor- communication, contamination Fig. 2013). on 1, Since there the the is fungal no spores indication (D. of Horne, UBC Eurotiomycetes and Flanagan (1982). Comparing just the those from Jayaweera andbrevicompactum Flanagan (1982) suggests that:and (2) (1) even within the same genus, the to measure ice nucleationThe ability results from of Jayaweera and spores Flanagan(filled from (1982) symbols). are the The included results three in from classes Fig.they Pummer 3 did investigated et for here. not al. comparison (2013) reportcalculate are the this not cumulative value included from in number their Fig. of experimental 3 IN description. since per sporeice and nuclei it per spore was is not equal possible to to 0.01, all of the Also, on average, themycetes order ofthe ice cumulative nucleating number ability ofthe ice for nuclei these per sporesspores, spore and is was 0.01 0.01 at approximately at approximately and 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | C S. ◦ er- ◦ Eu- ff 35 ects of − ff T > ≥ cient ice nuclei 25 ffi − values for the s n erence in surface area ff spores were assumed to spectra for all of the fungal s n , for the fungal spores in the ), we approximated all spores s values for Asian mineral dust n aer s spores. The A n C, but unity for ◦ 25 − P. frequentens T > erence in surface area per particle, we cal- 5030 5029 and ff Eurotiomycetes S. This can be explained by increased downward ◦ . Finally, Fig. 4 shows that, at temperatures below 100 %). Positive non-zero values indicate increased values for the fungal spores studied here are shown s × n spores were approximated as spherical with diameters P. notatum spores to be prolate spheroids having length and width C, whereas negative non-zero values indicate decreased ◦ Penicillium 35 − erence in freezing temperatures within the same genus observed C); in the second scenario (IN-Active), ice nucleation scaveng- ◦ ff C, all of the fungal spores studied here are less e ◦ . (2) Aspergillus , was approximated to be a prolate spheroid having dimensions 35 20 − erences between simulations with 3 µm and 8 µm particles are minor. Aspergillus ff C and − ◦ spores studied by Jayaweera and Flanagan (1982) were calculated using aer 25 5.5 µm; and both A − × P. digitatum INperSpore = Figure 5 illustrates that the inclusion of ice nucleation scavenging of fungal spores After accounting for surface area, Fig. 4 shows that the The temperature dependent s This explanation is supported by results shown in Fig. 6, which shows the percent spores over the oceans andsimulations in was polar land regions. Since surfaces,centrations the of excluding source fungal land of spores ice, fungal overlong-distance these spores the transport regions in decrease can of the in be fungaldue explained the to spores by surface ice a from nucleation decrease con- land followed incleation by surfaces the in precipitation. to In mixed-phase these stark clouds contrast,the remote increased scavenging region regions by surface between ice concentrations roughly nu- transport of 50–80 of fungal ice-borne spores spores in titudes, from mixed-phase where clouds ice at evaporates higher and altitudes to releases lower spores al- in the region between 50–80 perature range. The results fromfigures, these the simulations top are and givenrespectively. bottom Di in panels Figs. correspond 5 to and results 6. for In 3 both µmin and mixed-phase 8 clouds µm can particles, decrease the surface annual mean mixing ratios of fungal (i.e. below 0 to ing for fungal spores(see Table was 2). Results set from the toence simulations zero in are presented the for in fungal termsActive spore of – percentage mixing IN-Inactive)/IN-Inactive di ratio betweenconcentrations the due two to scavenging scavenging scenarios frombetween (i.e. heterogeneous (IN- ice nucleation atconcentrations temperatures due to scavenging from heterogeneous ice nucleation over this tem- We used the globalice chemistry-climate nucleation transport on model theTwo scavenging EMAC scenarios transport to were and compared: study in distributionation the the scavenging first of e by scenario fungal fungal (IN-Inactive), spores ice spores was nucle- set in to zero the for atmosphere. mixed-phase cloud temperatures 3.5 Modeling results 7.75µm be spheres with a diameter of 3.25 µm. species studied here are similar.cannot Figure explain the 4 di also showsin that Fig. a di 3 forapproximately species of compared to Asian mineral dust on a per surface area basis. (Niemand et al., 2012)Jayaweera (experiments and ACI04_16 Flanagan and (1982) ACI04_19), for rotiomycetes and also results from spore dimensions and morphologiesically, from the literature (Raper et al., 1949). Specif- n To calculate the surface areaother for each than fungal spore the ( shown in Table 1. given in Table 1. in Fig. 4. For comparison, included in Fig. 4 are culated the ice nucleationcurrent active study. The surface ice site nucleation active density, of surface ice site density nucleation corresponds sites to2009), per the and unit number was surface calculated area according of to: a particular type of IN (Connolly et al., nucleation. To take into account the di 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | P. P. P. IN- sp., . The Ascomy- Basidiomy- ) cannot be spores in re- Puccinia 17.5 µm for fungal spores. 25.0 µm for spores are not Basidiomycota P. triticina × × 3.2 µm (based on spores are not all × ). Within both phyla , this spore was ap- ) are the poorest IN, s n 6.0 µm (Schubert et al., Ascomycota , 28.3µm × 20.6 µm for 20.4 µm for Ascomycota × erent spore species. Finally, Basidiomycota × spores being better IN com- and ff S, there is an increase in con- Basidiomycota ◦ and C. herbarum Basidiomycota spore studied by Jayaweera and , P. striiformis , 25.0µm P. brevicompactum , and to calculate sp. studied by Iannone et al. (2011), we and 23.3µm together make up 98 % of known fungal sp. from Iannone et al. (2011), the spores Basidiomycota Basidiomycota 5032 5031 for fungal spores from the classes spores relative to Ascomycota Basidiomycota for the spores studied by Morris et al. (2013b), spores. Figure 8 also shows that the variability sp. and s A. brasiliensis 19.7 µm for s n , × n spores. Figure 7 also shows that the ice nucleating spores studied by Jayaweera and Flanagan (1982) erence in the ice nucleating ability of the ff ) there is a wide range of freezing properties. Also, the P. lagenophorae H. vastratrix, Ascomycota Cladosporium fungi, we have compared the cumulative number of ice . The other for Penicillium Cladosporium , 24.4µm Ascomycota and Ascomycota 3.4 Penicillium for Basidiomycota spores (Fröhlich-Nowoisky et al., 2012). To investigate further P. frequentens Ascomycota erence in surface area between di s Cladosporium herbarum . To calculate P. allii ff , n 12.5 µm for 23.0 µm for vs. spores ( × × Basidiomycota fungal spores in Figs. 7 and 8, respectively. The data are taken from for the s INperSpore n Basidiomycota and Basidiomycota P. notatum values shown in Fig. 7 were provided in the original studies (Haga et al., 21.7 µm for Ascomycota Ascomycota , , 14.6µm , 32.0µm × Ascomycota Ascomycota and the and Ascomycota Figure 8 shows that, once normalized to surface area, Figure 8 shows previous results of To calculate Ascomycota was discussed in Sect. ically, we used dimensionsaristidae of length25.0µm and width asgraminis follows: 29.0µm calculation of better IN compared to ability of Asian mineral dust particlesin studied the by middle Niemand of et the al. range (2012) observed is for approximately cota we approximated the spores asmorphology prolate spheroids and based size onAnikster literature ranges et values for (Cummins al., spore 2005; and Anikster Husain, et 1966; al., Bruckart 2004; Laundon et and al., Waterston, 2007; 1964). Specif- but on the otherdigitatum hand, some of( the spores from thisvariation within phylum each are phylum is also greaterproperties the than of the best each variation IN phylum. between Overall, the ( we average conclude freezing that of the perSpore 2013; Morris et al., 2013b; Jayaweera and Flanagan, 1982). Based on Fig. 7, some cota nuclei per spore andand the ice nucleationthe active current surface study site aset density al. well of (2011), as Jayaweera and from Flanagan the (1982), and following Morris studies: etreanalysed Haga al. the (2013b). et results al. fromscribed (2013), Iannone above Iannone et (i.e. al. procedures (2011) described using in the Sects. same 2.2 procedure and as 2.3). de- All of the other species found on Earthcreased concentrations (James of etmote al., marine regions 2006). and Recent it wasthat field suggested (based these measurements on preliminary findings found ice may nucleationpared in- results) be to explained by whether there is a systematic di centrations of fungal spores when iceexplanation for nucleation this in observation mixed-phase clouds is is thatfollowed included. ice by An nucleation and evaporation precipitation of atincreased high the fungal altitudes, spore ice concentrations crystals at these at altitudes. heights3.6 between 0 and 5 km, leads to The phyla lustrates that the inclusion of icea nucleation decrease scavenging in in mixed-phase mean cloudsmid-latitudes, mixing leads ratios where to in precipitation the scavengingcontrol mid- in on to mixed-phase aerosol upper clouds transport troposphere,2012). particularly to exerts Below polar 5 in a km regions the strong and (Bourgeois in the and region Bey, between 2011; 50 Zhang and 80 et al., change in zonally averaged vertical profile of annual mean mixing ratio. This figure il- all better IN comparedin to freezing properties withinaccounted for both by phyla a ( di proximated to be a prolate2007). spheroid with To dimensions calculate 19.5µm were approximated to be prolatemicroscope spheroids images with dimensions of 4.6µm the spores Iannone et al., 2011). Flanagan (1982) was 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | IN- . We C, and ◦ . On a per 31 − spores. C and ect of IN activity on cient cloud conden- ◦ Eurotiomycetes ff ffi ects. ff 25.5 Ascomycota Eurotiomycetes , and − ' C). We used our new freezing , which together make up 98 % ◦ and Bertram, 2010; Cziczo et al., C, all of the fungal spores studied ff erent species of fungal spores cho- ◦ ff C, all fungal spores for which data are 20 ◦ − ) there is a wide range of freezing proper- 20 ect of acidic coatings on the ice nucleation Ascomycota ff − Agaricomycetes Ustilaginomycetes C, 0.01 between ◦ 5034 5033 , > and 29 − spores are better IN than C (Fig. 3). On average, the order of ice nucleating ability Basidiomycota ◦ C and ◦ 36 19 and − Agaricomycetes − Basidiomycota Ustilaginomycetes Basidiomycota cient ice nuclei compared to Asian mineral dust on a per surface C and ◦ ffi 26 − Ascomycota value of 0.01 can vary by approximately 20 In the simulations we assumed that the fungal spores were e To our knowledge, this study is the first to model globally the e Comparing our results with previous results, we show that within the same genus, All of the spores studied caused freezing of water droplets above homogeneous itz et al., 2011; Salam2008). et al., 2007; Sullivan et al., 2010; Yang et al., 2011; Zobrist et al., mersion freezing only. Experiments thatin explore other ice freezing nucleation modes bysults. these would Also, fungal be experiments spores useful, that and exploreability the they e of would complement these thethe spores current ice would re- nucleation ability beArchuleta of useful the et as spores, al., similar 2005; these2009; to Attard coatings Gallavardin mineral et et dust could al., al., (Eastwood 2008; 2012; potentially2010; et Kanji Cherno Koop et al., change and al., 2009; Zobrist, 2008; 2009; Knopf Möhler and et Koop, al., 2006; 2008; Koehler et Niedermeier al., et al., 2011, 2010; Re- While the treatment of iceindication nucleation of scavenging the is regions simplified, that this areby study most ice gives sensitive an nucleation, to initial as removal well of spores as due the to potential scavenging magnitude ofsation these nuclei, e as donethis previously assumption (Heald are and needed. In Spracklen, addition, 2009). the Experiments freezing results to shown verify here were from im- fungal spores studied canby modify providing their atmospheric an transport additionalSpecifically, we and show removal global that mechanism the distributions inclusion ofmixed-phase of clouds ice the can nucleation (1) spores scavenging of decreasespores from fungal the spores over surface the in the annual atmosphere. mean oceans mixingratios and in ratios polar of the regions upper fungal troposphere. and (2) decrease thethe annual mean distribution mixing of fungal spores in the atmosphere and their long-distance transport. ties and that not all 4.2 Global transport of fungal spores Using a global chemistry-climate transport model, we show that ice nucleation on the both phyla ( of known fungal species found on Earth (James et al., 2006). The data show that within (Fig. 4). In addition,here were at less temperatures e below area basis. the freezing temperatures can varyperSpore drastically (e.g. the freezingresults temperature together at with an dataspores from from the the literature phyla to compare the freezing temperatures of freezing temperatures. The cumulativetemperatures number between of ice0.1 nuclei between per sporefor was these 0.001 spores was at surface area basis, all of the fungal spores studied had similar freezing properties 4.1 Ice nucleation on fungal spores To add to the limitedwe amount studied of the data on icesen the nucleation from ice properties nucleation three of properties classes: 12 offocused di fungal on spores, these classesand since also they because are thereof is thought spores currently to from little be these quantitative classes. abundant data in on the the ice atmosphere, nucleation ability available are poorer immersion IN than Asian mineral dust on a per surface area4 basis. 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Z., Braun, U., Dijksterhuis, J., Starink, M., Hill, C. F., Zalar, P., de Shelton, B. G., Kirkland, K. H., Flanders, W. D., and Morris, G. K.: Profiles of airborne fungi in 5 5 30 25 20 15 10 30 20 25 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , , Spores/Drop 10.1021/jp7112208 spores with an uncertainty of 10.1007/s10453-006-9022-2 2) basidiospore 28 2) basidiospore 4 , resulting in uncertainties of approximately (µm) Spore Type Median 2) basidiospore 3 2) basidiospore 6 2) basidiospore 15 2) teliospore 9 2) teliospore 4 2) teliospore 15 1) conidiospore 15 1) conidiospore 6 ± ± ∗ ± ± ± ± ± ± ± ± 7( 6( , 2012. Ustilaginomycetes 7( 5( 4( 6( 7( 6( 3( 3( × × × × × × × × × × 2) 2) 2) 2) 2) 2) 2) 2) 1) conidiospore 2 1) conidiospore 3 1) 1) ± ± ± ± ± ± ± ± ± ± ± ± and the Eurotiomycetes 8( 10( 13( 8( 7( 7( 9( 8( 4( 4( 4( 5050 5049 Agaricomycetes were used for the × sp. 4( 10.5194/acp-12-8911-2012 , and 50 × , 40 × Amanita muscaria Boletus zelleri Lepista nuda Trichaptum abietinum Ustilago nigra Ustilago avenae Aspergillus niger Penicillium Penicillium brevicompactum objective was used for the × Description of the fungal spores investigated in the current study. tions: the role of water activity, J. Phys. Chem. A, 112, 3965–3975, doi: spores prevalent in the aerosola of 2-year the period city (2001–2002), of2006. Aerobiologia, Caxias 22, do 119–126, Sul, Rio doi: Grande do Sul, Brazil, over HAM, version 2:Phys., 12, sensitivity 8911–8949, doi: to improvements in process representations, Atmos.2008. Chem. ClassAgaricomycetes Agaricus bisporus Species Spore Size Ustilaginomycetes Ustilago nuda Eurotiomycetes Aspergillus brasiliensis The uncertainty in the size measurements correspond to the resolution limit of the Olympus IX70 and Zeiss Axiolab ∗ A microscopes. The 20 approximately 2 µm, and the 30 1 µm. Table 1. Zobrist, B., Marcolli, C., Peter, T., and Koop, T.: Heterogeneous ice nucleation in aqueous solu- Zoppas, B. C. D., Valencia-Barrera, R. M., Duso, S. M. V., and Fernández-González, D.: Fungal 5 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | B. , ), and ) C) ◦ 35 nuc, ice R A. muscaria U. avenae ≤ − , ). T evicompactum , and . br P U. avenaeU. L. nuda L. 35 C) ( ◦ − A. bisporus 35 ( U. nigra T > − , 25 0.05 ≥ − 25 > T > T > − P. brevicompactum C U. nuda ◦ ( sp., (0 temperature range 1 for ) Mixed-phase clouds Ice clouds Agaricomycetes B. zelleri U. nigra U. Penicillium sp. 5052 5051 nuc, liq R Penicillium , Ustilaginomycetes ), A. niger , scavenging ( A. niger U. nuda U. A. muscaria T. abietinum A. brasiliensis ( , and Nucleation scavenging schemes used in EMAC. SEM images of spores from the classes L. nuda Scavenging scheme Liquid nucleation Ice nucleationIN-Inactive scavenging ( IN-Active 1 1 0 for entire 0 for 0.05 , . abietinum A. brasiliensis A. bisporus T Fig. 1. zelleri Eurotiomycetes Table 2. Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , ) A. Eu- spores : ] m 2 spores ( Ustilagino- ) are in red u 8 t ) are in red 9 : A. bisporus c . ] 1 s s a

p 1 , i u s p s 1 n

s o m a INperSpore(T) s n a 0 m i e e m u t e g u r 2 e o s i n u n i e s r [ a l a c a i i

e e r a i i c e r l o t . n t a c n t a Ustilaginomycetes l r s l e g v y e e s e a p i d l e e d i l g a e a o l s g c u c

u i v m c : i r i r m u b i e F t spores ( y y

m n n a o y u b m z b n b n a n e

t

d

o . . . m n d . . . . . e . . . m c i e n o u s o a H A A B L T U U U A A P P s g t a n Eurotiomycetes c i

i

n p s i t a s o

m e a l r ) (Iannone et al., 2011), m o

t n m s i n r u

a r i t u e m n o e t n x n t i s g u h e s i c u e a l s a a t i t i i r u u t u Agaricomycetes I T A U E : w r e s v i a e e r ] q i t d ) for e . a a a g

a g o e 2 i b a r i o l r i y c p a r r n P. brevicompactum p 1 r a a s b d n f s b n a a d

s e e

r P. brevicompactum s 0 g i u l J d u i v s s e l e 2 [ m m m m m m b t

u n n a u u m e n [

s u 0 l l u u u u u

i e s s z s n t . l l i i i i i e

o o o c e e a 1 l i i l l l l l t u p t t t s sp., m l l l l l - y a g g g e a i g g c i i i i i e e t a u

i a a a r r sp., c n t c c c c c n t s m c c : h l l l r i i i i i i e e y e i i i a e a y y o c y

a l t t t i n n n n n p p p i Agaricomycetes m n s s s s d d g m o s s e e e e e m m e r i o u n o o A A B L T U U U A A P P P P P A g t c i i

a s i t t a

l r ) are in blue (open symbols); o o 5 s

i m i a t r r e 1 s i h g u u - T A U E E N INperSpore(T) ) Penicillium C o 0 , ( Penicillium 1

- 0 e , ) are in blue (open symbols); 2 r ) are in green (open symbols with horizontal line); - u 5054 5053 t a 5 r T. abietinum 1 e - , p

A. niger m 5 , ) are in green (open symbols with horizontal line); e A. niger 2 ) - T 0 ,

C 2 o - g ( ) for submicron Asian dust studied by Niemand et al. (2012) U. avenae

n e , i r L. nuda T. abietinum z u t e

, , a 0 5 e r r 2 3 e - - F p m U. avenae e , U. nigra T 0 , 3 A. brasiliensis - L. nuda 5 B. zelleri 3 , A. brasiliensis - , 5 3 INperParticle(T) - U. nigra U. nuda , spores ( 0 4 - 0 0 3 1 2 spores ( B. zelleri - - - 4 - 0 0 0 0 2 3 4 1 0 - - - - , 1

0 1 1 1 0 0 0 0

1

1 1 1 1 z o r F s t e l p o r D f o n o i t c a r F n e r o e r o p S r e p N I # e v i t a l u m u C e l c i t r a P A. muscaria spores ( , U. nuda Fraction of droplets frozen as a function of temperature. Cumulative number of ice nuclei per spore as a function of temperature ( Fig. 3. A. muscaria as derived from the measurement data shown in Fig. 2. Fig. 2. (open symbols withobtained vertical using line). the same Homogeneous apparatus freezing as this results study, are ( also included. mycetes Eurotiomycetes spores ( rotiomycetes (open symbols with vertical line). Also shown is ( studied by Jayaweera and Flanagannuclei (1982) per (filled particle symbols), ( and(half-filled the diamonds). cumulative number of ice bisporus Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ) T. A. , , : ] 2 U. avenae L. nuda 8 , 9 , 1

, n a g a A. brasiliensis n a l U. nigra m F ,

u t B. zelleri d c n s a , s a i

n p s e m a m t n m r u u m n e o e t spores ( n t i e u s i c e a l s a t t i i i r u C. Constant emissions were u t u : w r e s v a i e r ◦ U. nuda ] q i t d e . e a a a g

g e o 2 i b o a r i l a r i y c p a r r p 1 r a a n f s s b d n b n a a d

s e

r

s 0 g i u l e J 35 d u i s s e l e [ 2 m v m m m m m b t

u n n u u m e n [

s u a u u l l u u u −

i e s s z s n t . i i l l i i i e

o o c e e a l l l i i l l l t u p o t t t s m l l l l l y a g g e a

i g g i i c i i i e e t a g u

i a a r r A. muscaria c c c n n t c c c t s m c c : h l l a r i i i i i i e e y a e i i l e a y y spores ( o c y

i a l t t i n n n n n , p p p i ), for fungal spores determined using m n s s s s d d e e g m o s s e e e m m e r i 2 o u n o o P P A A A B L T U U U A A P P P g t c i i

− C and a s i t t a

◦ l

r o o s i m i a t r r e s (m i h g u u Eurotiomycetes T A U E E N 25 s − n 0 1 - A. bisporus 5056 5055 5 ) are in red (open symbols with vertical line). Also 1 - Ustilaginomycetes 0 spores ( ) 2 - C o (

100 %. Top: 3 µm particles; bottom: 8 µm particles. In all sim- e r u

× t 5 a 2 r - e p m e T 0 3 - P. brevicompactum 5 Agaricomycetes 3 - sp., 0 values for Asian mineral dust (Niemand et al., 2012) (half-filled diamonds). 4 - 7 6 5 9 8 0 1 s 0 0 0 0 0

0

1 1 1 1 1

n

1 s

) m (

n ) are in blue (open symbols);

2 - Ice nucleation active surface site density, Global model results showing percent change between IN-Active and IN-Inactive EMAC Penicillium , Fig. 5. simulations in surface annual meanActive – mixing IN-Inactive)/IN-Inactive ratio, with percentulations, change spores being are calculated assumed to asby be (IN- ice-phase CCN-active. nucleation IN-Active at temperatures assumes between that particles are removed simulated from land surfaces except for land covered with ice. the data in Fig. 3. Fig. 4. abietinum are in greenniger (open symbols with horizontal line); shown are Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | C. ◦ 35 − ] 2 1 0 2

, . l a

t e C and

d ◦ m n : i i ] u a s t

2 s c m 8 e m s a e s e 9 i i n u 25 n p a s 1 r k m e N r [ m t r n m a [

u s o

u m e n a x o b i − t n n t i i h s s e s i e u r c h a l s a i i a r t t i . i m i p a a r e u u a t t u g e r n r n s v i a p e e r o n n : h q i i d i m a t d i i e .

s a a o a ] i g a g o n e r : i b f u a t r i o c c r i l c p t a n m m i i 1 i r i ] e n s i r p i i m m r s : a s i i a i b d n f s t s i b n t t a d a a i . a

s e c 1 e 3

l g l ]

r u u g i r i r l i u v i l a r l r l r i i d u i v p t s s 0 1 e r t a r 3 l m a e v m m m m m b r r F g t a i a l s a s g t u n n a u u m e 2

0 n 1

u n a r

l l u u u u u o o i [ z s n t l t a l l i i i i i 2 0 d a a a a a a a a a a

o o o o a a i p p i i l l l l l . i i i i i i i i i i [ u p t t s 2 r m n l l l l l l

s s a g g g e a a i g g

: c n n n n n n n n n n i i i i i . [ t a l o c u i i a i a i i i i i i i i i i

l o o a a a r r i n t t t c c c c c n

a s

. c h l l l : : r o c c c c c c c c c c i i i i i t i e e t a d d l e e e i i i a a a y c

y y m a l c c c d l l c c c c c c c t t t i n n n n n e o values for submicron Asian p p p i a a a t r l l

s s s e s

d d l l g m o u u u n u u u u u u u s s e e e e e c e r i i m e e t e

u u y o e A A B L T U U U P P P E T T P P P P P P H P A A P P P P P C C A e t t s n i

i s s m w o d

a r i o a s s n r g s i i c y n o a s a h h a a J I H M A T B T

spores investigated in this study

100 %. Top: 3 µm particles; bottom: × 5 - INperParticle(T) 0 1 - Basidiomycota 5058 5057 5 and 1 - ) C o (

e r 0 u t 2 - a r

e p m e T 5 2 - Ascomycota ) for 0 3 - 5 3 - 5 1 2 3 4 0 - - - - - 0 0 0 0 0 0 1

1 1 1 1 1

INperSpore(T) e l c i t r a P r o e r o p S r e p N I # e v i t a l u m u C Global model results showing percent change between IN-Active and IN-Inactive EMAC Comparison between the cumulative number of ice nuclei per spore as a function of tem- mineral dust (Niemand et al., 2012). and earlier studies including Haga et(1982), al. and (2013), Morris Iannone et et al. al. (2011), (2013b). Jayaweera and Also Flanagan shown are perature ( Fig. 7. Emissions were simulated from land surfaces except for land covered with ice. 8 µm particles. In all simulationsparticles spores are are assumed removed to by be CCN-active. ice-phase IN-Active assumes nucleation that at temperatures between Fig. 6. simulations in zonally averaged vertical profilebeing of annual calculated mean as mixing ratio, (IN-Active with percent – change IN-Inactive)/IN-Inactive Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ) for T ] vs. 2 1 0 s 2

, . n l a

t e

d m n : i i ] u a s t 2 s c m 8 e m s a e s e 9 i i n u n p a s 1 r k m e N r [ m t r n m a [

u s o

u m e n a x o b i n t n t i i h s s e s i e u r c h . a l s a i i a r t t i i m i p a a r e u u a t t p u g e r n r n s v i a e r o n n : h s q i i d i m a t d e i i . e

a a o a ] i g g o n e r : i b f u a t r i o a c c r i l c p t n a m m i i 1 i r i ] e s i r p i i m m r s : a s i i a i s b d n f t s n i b n t t a a d a i . a

s c 1 e 3

l g l ]

r u u g i r i r l i u e v i l a r l r l r i i d u i p t s s 0 1 e r t a r 3 l m a v e v m m m m m b r r F g t a i a l s a s g t u n n u u m e 2

0 n 1

u n : a a r

l l u u u u u o o i [ z s n t l t a l l i i i i i 2 0 d a a a a a a a a a a

o o o a a i p p i i l l l l l . i i i i i i i i i i [ u p o t t s 2 n r m l l l l l l

s s a g g e a a i g g

: c n n n n n n n n n n i i i i i . [ t a g o l c u i i a a i i i i i i i i i i i l

o o a a r r i n t t t c c c c c a n s

. c : : h l l a r o c c c c c c c c c c i i i i i t i e e a t l d d e e e i i l a a a y

c y y m a l c c c d l l c c c c c c c t t i i n n n n n e o p p p a t i a a r l l

s s s e s

d d l l g m o u u u n u u u u u u u s s e e e e e c m e r i i e e t e

u u y o e A A B L T U U U P P P E T T P P P P P P H P A A P P P P P C C A e t t s n i

i s s m w o d

a r i o s a s n r g s i i c y n o a

s a h h a a

H M A T J I B T

0 5 - 0 1 - 5059 5 1 - ) C o (

e r u t 0

a 2 r - e p m fungal spores calculated using the data in Fig. 7 and using ap- e T 5 2 - 0 3 - 5 3 - Basidiomycota 0 4 and - 0 5 4 9 8 7 6 1 0 0 0 0 0 0

0

1 1 1 1 1 1

1

s

) m ( n

2 - Comparison between ice nucleation active surface site density spectra ( Ascomycota proximations for spore size andstudy shape detailed as in well the as text.Jayaweera Included and results are Flanagan from results (1982), from the and theous following Morris current results studies: et for Haga al. submicron (2013b). et Asian Also mineral al. shown dust (2013), in (Niemand Iannone the et et figure al., are 2012). al. previ- (2011), Fig. 8.