Review

Tansley review The occurrence of red and yellow autumn leaves explained by regional differences in insolation and temperature

Authors for correspondence: Susanne S. Renner1 and Constantin M. Zohner2 Susanne S. Renner 1 2 Tel: +49 89 17861 250 Systematic Botany and Mycology, University of Munich (LMU), Menzinger Str. 67, Munich 80638, Germany ; Institute of Email: [email protected] Integrative Biology, Department of Environmental Systems Science, ETH Zurich, Zurich 8092, Switzerland

Constantin M. Zohner Email: [email protected] Received: 19 January 2019 Accepted: 24 April 2019

Contents

Summary 1464 IV. The adaptive value of colour-changing leaves 1468

I. Introduction 1464 V. Outlook 1469

II. Phylogenetic and geographical occurrence of autumn colour Acknowledgements 1469 change 1465 References 1470 III. Physiological functions of autumnal leaf xanthophylls and anthocyanins 1466

Summary

New Phytologist (2019) 224: 1464–1471 Red or yellow autumn leaves have long fascinated biologists, but their geographical doi: 10.1111/nph.15900 concentration in trees in Eastern North America (ENA) has defied evolutionary explanations. In this review, anthocyanins and xanthophylls are discussed in relation to their occurrence in Key words: adaptive explanation, different regions of the Northern Hemisphere, phylogenetic distribution and photoprotective anthocyanins, photo-oxidative damage, function during the breakdown of chlorophylls. Pigments in senescing leaves that intercept regional climates, solar irradiation, incident light and dissipate the absorbed energy extend the time available for nutrient resorption. xanthophylls. Experiments with Arabidopsis have revealed greatest anthocyanin photoprotective function at low temperatures and high light intensities, and high-resolution solar irradiation maps reveal that ENA and Asia receive higher irradiation than does Europe. In addition, ENA experiences higher temperature fluctuations in autumn, resulting in cold snaps during leaf senescence. Under common garden conditions, chlorophyll degradation occurs earlier in ENA species than in their European and East Asian relatives. In combination, strong solar irradiation, temperature fluctuations and, on average, 3-wk shorter vegetation periods of ENA species favour investment in pigments to extend the time for nutrient resorption before abscission, explaining the higher frequency of coloured species in ENA compared to Europe. We end by outlining research that could test this new explanation of bright New England autumns.

followed by the eventual shedding of most leaves. During this I. Introduction process, plastid-located carotenoid pigments (xanthophylls), pre- In the temperate zone, decreasing day length and cold temperatures sent throughout the year but initially masked by Chl, become induce the breakdown of leaf chlorophyll (Chl) during senescence, visible. These pigments play a major role in deactivating the Chl

1464 New Phytologist (2019) 224: 1464–1471 Ó 2019 The Authors www.newphytologist.com New Phytologist Ó 2019 New Phytologist Trust New Phytologist Tansley review Review 1465

3 * 1 * triplet state ( Chl ) and singlet oxygen ( O2 ) formed during light (a) (b) excitation and help dissipate excess light energy (Ruban et al., 2002). The Chl breakdown process often is accompanied by the synthesis of vacuole-located anthocyanin pigments that also have a protective function against excess light and reactive oxygen species (ROS) (Chalker-Scott, 1999; Close & Beadle, 2003; Lee et al., 2003; Gould et al., 2018; Xu & Rothstein, 2018). During the past 20 yr, the possible adaptive value of yellow- or red-coloured autumn leaves has attracted the interest of both physiologist and evolutionary biologists. Indeed, Lev-Yadun & Holopainen (2009) mention that over 80 papers have discussed the adaptive value of autumn colouration. A review of autumn colour change published 10 yr ago, based on Fig. 1 Autumn leaves of the Asian species Cornus kousa (a) and the North- a meeting, ended on ‘the uncomfortable fact that there seem to be American Vaccinium corymbosum (b) showing that anthocyanin production more hypotheses, and more published reviews and opinion pieces, is sun-induced. Shaded parts of leaves remained yellow due to the lack of on the subject of red leaf colouration than sources of good anthocyanin, whereas sun-exposed parts turned red. Feild et al. (2001) experimental data’ (Ougham et al., 2008, p. 13). Over the past observed a similar pattern in Cornus stolonifera. Photos: E. I. Arndt. decade, better data have become available on the phylogenetic and geographical distribution of the phenomenon of autumn coloura- Holopainen, 2009). We end by laying out a new explanation for tion, the pathway of Chl degradation and the protection of why leaf colouration is more critical for North American and Asian senescing cells from oxidative damage (Archetti, 2009; Lev-Yadun woody species than for European species. We also outline further & Holopainen, 2009; Landi et al., 2015; Viola et al., 2016; Gould research that could clarify the adaptive value of red and yellow et al., 2018). These data prompted us to revisit the function and leaves, and the role played by regional temperature and irradiation evolution of autumn colouration. differences that affect the phenological behaviour of long-lived A key aspect of Chl breakdown is the recycling of nutrients, woody plants. especially the nitrogen (N)-rich proteins in the organellar cell walls (Matile et al., 1999; Keskitalo et al., 2005). Nutrient resorption II. Phylogenetic and geographical occurrence of needs a functioning metabolism, which in turn requires that cells be autumn colour change protected from oxidative damage when photons are no longer fully used in photosynthesis. This sets up the expectation that autumn Transplantation experiments have shown that individuals maintain colouration should be especially important under environmental their autumn colour change when grown in different places conditions of high light intensities and low temperatures, damaging (Archetti et al., 2009), and many predictably colour-changing senescing leaves (Fig. 1). This expectation can now be tested against cultivars are available commercially, implying a strong genetic basis ‘big data’ temperature and irradiation maps that provide high- for patterns of autumn colour change. Even so, before 2009, there resolution measurements on the long-term monthly averages of were no data on the phylogenetic distribution of autumn leaf temperatures and solar irradiation in North America, Europe and colours. In that year, Archetti (2009) compiled the autumn leaf Asia, the continental regions most relevant to the topic of autumn colours of 2368 tree species from 400 temperate zone genera, colouration in arborescent species. In combination, the new including both deciduous and evergreen groups, and then recon- climate and irradiation data, experiments on the environmental structed the evolutionary origin of red and yellow autumn leaves conditions under which anthocyanins are most beneficial for using the 567-taxon phylogeny of Soltis et al. (2007) and standard senescing leaves (Gould et al., 2018), the relative frequencies of red, parsimony reconstruction. Archetti categorized species as having yellow and noncoloured leaves in different regions of the World, (or not) green, yellow or red autumn leaves, and for a few also and common garden experiments on Chl breakdown in Northern distinguished yellow-orange, brown-yellow-orange, yellow-red Hemisphere species (Zohner & Renner, 2017; Zohner et al., 2017) and other combinations. He found red autumn leaves in 290 have led to a new framework for an adaptive understanding of (12%) species from 70 (17.5%) genera and yellow (independent autumn colouration. from red) in 378 (16%) species from 97 genera (24%). The list In this review, we begin with the geographical and phylogenetic includes hundreds of nondeciduous species (e.g. 125 species of distribution of autumn colour change, summarize current phys- Rhodendron,53ofPinus, numerous Cupressus, Cotoneaster) as well iological understanding of the function of anthocyanins and as arctic, temperate and subtropical species. xanthophylls in senescing leaves, and present the arguments in An analysis of autumn colour change in 1308 individuals from support of the photoprotection hypothesis (Gould et al., 1995, 711 deciduous woody species in a common garden in Munich 2018; Feild et al., 2001; Hoch et al., 2001; Wilkinson et al., 2002) (48°090N, 11°300E; 501 m asl) by a team of three observers from and those for bright ‘signalling’ colours as resulting from coevo- September to November 2017, found pure red autumn leaves in lution with herbivorous insects (Archetti, 2000; Hamilton & 5% (32) of the species, yellow and red leaves in 32% (228), and pure Brown, 2001; Archetti & Brown, 2004; Archetti et al., 2009), yellow leaves in 63% (451). There were no deciduous noncolour- including extinct herbivore–plant interactions (Lev-Yadun & changing species. In combination these data show that – as pointed

Ó 2019 The Authors New Phytologist (2019) 224: 1464–1471 New Phytologist Ó 2019 New Phytologist Trust www.newphytologist.com New 1466 Review Tansley review Phytologist

out by Archetti (2009, p. 707) – coloured senescing leaves are rare III. Physiological functions of autumnal leaf in flowering plants and gymnosperms overall, and it ‘should not be xanthophylls and anthocyanins taken for granted that they are a normal side-effect of leaf senescence’. However, in deciduous temperate-zone trees and Both young leaves and senescing leaves are at risk of photo- shrubs of the Northern Hemisphere they are common. This point is oxidation damage because their photosynthetic apparatuses are highlighted by comparing a phylogeny of 1369 temperate species incomplete and unable to deal with high irradiance (Juvany et al., including deciduous and evergreen species (Fig. 2a) and one of 501 2013). In young growing leaves, damage can occur because of the deciduous-only species (Fig. 2b), with the autumn colours of their intense generation of ROS due to low ascorbate peroxidase activity. leaves colour-coded. In old leaves, it occurs because the first step in senescence is a rapid It has long been noticed that anthocyanin-coloured red autumn increase of ROS following the degradation of Chl, which leads to leaves are commoner in Eastern North America (ENA) than in the oxidation of pigments, proteins and lipids. These oxidation Europe or Asia, either because more species there turn red or (also) processes are needed for nutrient remobilization, but they are because red-leaved species are especially abundant (Lee et al., 2003; necessarily accompanied by a risk of oxidative damage (Introduc- Lev-Yadun & Holopainen, 2009). To quantify the phenomenon, tion). This is where autumnal pigments appear to play a role. Lev-Yadun & Holopainen (2009) examined the geographical Xanthophylls are the pigments responsible for the yellow-orange distribution of all of the species with red autumn leaves listed by colours of autumn leaves (Figs 1, 4). They are accessory light- Archetti (2009) and found that ENA has 89 such species, whereas harvesting pigments and present year-round in the photosynthetic the whole of Europe has only 24, a number that agrees with our own membrane or in the hydrophobic environment of plastoglobules compilation (Fig. 3). In Asia, at least 152 species have yellow or red (Matile et al., 1999). During leaf senescence and Chl breakdown, leaves (Lev-Yadun & Holopainen, 2009). This needs to be seen in xanthophyll concentration steadily declines in both yellow- and the context of the different tree species richness of these regions red-senescing leaves (Lee et al., 2003), and their role may be limited (Fine & Ree, 2006): The ENA temperate biome has 300 tree to protecting chloroplasts from short-term exposure to light stress species (at least 30% of them with red leaves), the European (Ruban et al., 2002; Gould et al., 2018). Anthocyanins are the temperate biome 124 (at least 19% of them with red leaves), and the pigments responsible for colours ranging from blue to purple and East Asian temperate biome 729 (at least 21% with red or yellow red, and sometimes also yellow (Figs 1, 4). They are highly water- leaves). soluble and are a cationic sub-class of flavonoids, with their precise Of 711 species studied in the common garden in Munich, 119 colour depending on vacuole pH and co-pigmentation processes came from Europe (i.e. 96% of Europe’s native tree species), 148 with other flavonoids, hydroxycinnamic acids or metallic cations. from North America (49% of the region’s tree species) and 444 Different from the xanthophylls, the anthocyanins’ role in from Asia (61% of the region’s tree species), and in this sample, photoprotection is becoming well-understood, at least mechanis- North America has a significantly higher proportion of species with tically as we describe in the following sections. red autumnal colouration than do Europe or Asia (P < 0.05; In the photo-oxidative context, anthocyanins in vacuoles (and Fig. 3). Of the North American species, about equal numbers turn other phenolic compounds in apoplast compartments) may have an red (47%) or yellow (53%). Europe again has the lowest proportion important role in scavenging ROS (Zhang et al., 2012; Xu & of species with red-coloured autumn leaves (20%), followed by Asia Rothstein, 2018), and in some cases anthocyanin biosynthesis is (38%). The colour change in all these deciduous species was under redox control (Page et al., 2012; Awad et al., 2015; Viola documented with photographs taken, on average, every 10 d per et al., 2016). Oxidative conditions induce anthocyanin synthesis individual, and the observer team agreed on the categorization of via redox-sensitive-transcription factors (Viola et al., 2016). Par- colours. Following Archetti (2009), orange was grouped with ticularly important is TCP15, a transcription factor that negatively yellow because both colours are caused by xanthophylls. The regulates anthocyanin accumulation. It is inactivated by oxidation phylogenetic distribution of the species with red- or yellow- under high-light conditions, and it always precedes anthocyanin coloured senescing leaves shows that red-coloured leaves occur in accumulation. In TCP15-overexpressing plants, the transcription 77 of 179 genera and are relatively evenly spread among temperate of PAP1, TT8 and DFR is lowered compared to wild-type (WT) zone clades (Fig. 2b). plants. Furthermore, ascorbate peroxidase-deficient mutants In St Louis, MO, USA, colour-change in marked red maple (apx1/tapx) and catalase-deficient mutants (Cat2HP1, Vander- (Acer rubrum) individuals has been monitored by the first auwera et al., 2005) accumulate more anthocyanin in response to author (2007–2018), confirming that cold sunny days intensify low temperatures (apx1 mutant; Asai et al., 2004) and to high light anthocyanin production (Lee et al., 2003; Fig. 4). This is (Awad et al., 2015). Put differently, anthocyanin biosynthesis genes supported by experimental cooling of autumnal maple are impaired by high light in peroxisomal catalase-deficient plants branches, which led to increased foliar sucrose, glucose and (Vanderauwera et al., 2005), and inhibition of catalase transcrip- fructose concentrations (two- to nearly 10-fold) and intensified tion after a cold night leads to increased photo-oxidation and anthocyanin production (Schaberg et al., 2017). In Arabidopsis, favours anthocyanin accumulation (Macrae & Ferguson, 2006). cold conditions activate the transcription of arogenate deshy- Comparative physiological experiments on the function of dratase, the enzyme that transforms arogenate into L-pheny- anthocyanins were first carried out in two unrelated species of lalanine, which also increases anthocyanin accumulation (Chen tropical understorey herbs (Begonia and Triolena) that each have et al., 2016). green and red leaves: Anthocyanins intercepted quanta otherwise

New Phytologist (2019) 224: 1464–1471 Ó 2019 The Authors www.newphytologist.com New Phytologist Ó 2019 New Phytologist Trust New Phytologist Tansley review Review 1467

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nu n n looi rrn nan a P in u s coc izzo a a F ini in a pl u y o a s P i u s o o g x xi s inin ia is n je u egieg e a s ld o s a P in uus s p f l nni n raxr u us x ox ldi n P s rir o f te icca i s F FraxFra in in a uus o i n a in u i re i i r in b gginose tal tta P i u s p g n ri a s x x F nthx e u n e a is i n u id ded yyi a anthoxa iie i petalp la v s P n u s ra e i ra x a s h i ulu a P i usu s m nng a r Fr FFrax rra lan c did g llusu ia P n d g o s F s u l l s P in uus s p u i e s u s s n s y iis I iin s t a a s u u us aan hhy ifolfo s a K l n u e rri t nns a inusnu inuin r s p tit li s K li u s g o tu ic a s x x innu inin innus d usu o ScS c oc a a x x aad h o la ensen fo u s a iuiu s p e rere c la raax ra x t ror n tiif u s S c d l oote ta aax a a u n e a us s T h surs m li g t F FrF r q tet ea s icc e a chc a o g e Fr FrFraF ant e rre u et nin d M r h is ur h llu t i s h y nnag r i i s ta i a i a u t i u mma ll unnau g o io P o c s a e s ii n s s i y n s rrgi i ata eei ii S s l h 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s ru uss e nnt lii lix p ne um P nu pep e iiffolo SaSali m u nns P ruun s m st y rpurp sii nusu toome ustu ra lix Sa rssi u s PPr u s t ng ife r li in rere Prr nnusu a ssi osos x if a P u s ra a m rere ollia r usu ce osa PPo S a pep ia P unun inos opu a rrii ensn r us ce p p lixlix nini s PPr nun s um i HyHyp HypeH Idde ul cac ffoo ru us vviumi ki p y e us ap lia PPr n a c erie p siaia tret pr ru s aa ricuc err p re e PrunusP nuu spin a a u icu o mum a n s m is ella FFllueggu m m lyyc ulal Pru u ncnc irrttell egg accm h ca a unun s i hhi E mos irirc rpa PPrru u ub ataata uuoo eea os cinun unusun s uull iaa nymusny a ssu epalep um r us rrr arri na mu u al m P nun serse pa y a Euo s fffrur um ruu s cu tatagg rpa e uttii PrP nu au n caca nnyymu ur coos ru s pe soo Eu EEuu mu oppa sa P bu us p rysry ttaa ono o s ae or g ch lla nymy nnyy m u SoS tataee s c eel EuoEu mu muus aaacka s ra gu ab ica Euouon on s v s ck CraC aee s flafl an n ymuym ve aallata iiii raratat gugu rmman illisis ymym u rrrru tu C ae ge imm uuss hams s ccoos s ratta s rsis lia E h em susu CCra iluilu pere oliao uo am eeno s sp uss alnif Eu nnymy iltoniton no Me eggu r on musu i vviii Mes taea hierhie liiaa ymum s ananus ra ncc tifofo Ceelas us o latit us CrC laan bu lis lastr oxyx ifool melme ar nnaal tru ypph liiuuss A ia mi s oor hyyl roonn ttoor a A rbiiccu llus ArA us llllossa ceerr ulatla oorrbbus vvii ga ca tuuss S nia longon A AcA ppililli hootitin obobl a ceerr ce ppee PPh nia ioos te r da s doniao ririaa ppeecic sisis A gm vid Cydy s a s s yen cer ent ii rbrbu ele tha mimicc osuosu SSoo om cacathayen AAccere rranan m aeenn eelles r d thhuu ChCha oomm A iiababolo m en sis A ceer t liiccuu Chhaea oensoen kii ceerr r tat m C s i nonossk cap arariicc alalusu cchoho AAc paadod um M s tsts cerer occiicu alu sca ppllaata cum M s fuus liliss AAcce tannoo alluu nttaa er ca ide MaM orieien s mmpp s alluus essttrisi AcAc essttrr MMa sylvlv er m e lus na Aceerr iyaiyabeb MaMa allia s cissifoiss ei lus hah enssii Ac ifoliuliu Malusa upehenh er velue m us hupeh A lutin Malul cacata cecer he umum s babac heldreildr alu oolldii AAcecer eichiihi MMa siebsieb sacacc i alulus nda haru MMa orriibundu Ac AAcecer m lusus ffll er momo opopal MMaa mmiila nspe us us pu ineanea ssusulanan MMaal a cocccic Acerc umum ccaannthah ana AcAcer er grigri PPyrayra orrtuneanane er pseudoe seuseum nthaha ffo udoppll yyraraccaant Acce atanutanus P rasasterer err cacarprpinif rusus pypyr inifooliliuum Py munis Accerer ssaca m us comom cchcharinuinu Pyr m s ppasashhiaia Accerer ruub Pyruus siiss bruumm ussurieienn AAccer ne PyPyruuss gundodo ifoifoliliaa AcAcer s PPyyrus pyr picatum rryayana Acer siebolie yrus ccaalllle boldidianumanum P lia A rusus betulbetulifolifo cer ppalalmaattuu PyPyr tuuss m nneaseastterer bbulullaat AAeesculuuss CoCototo flava ster mumultiflorus Aescu Cotonea lus glabrabraa dpressssus A Cotoneaster a esculus paparviflora s Cotoneastoneasterer foveoloveolatusu AAesescuullus hippocpocastanum Cotoonneeaaster aaccuuttifoolliuuss Koelreutteeria paniculata Cotoneaster dielsianus Phellodendroonn ssaacchhalinense Cotoneaster acuminatus Phellodendroellodendronn amamurenrensse Ginkgo biloba Tetradiumradium danielliidaniellii Metatasequoia glyptostrotroboidideess Zanthoxxylylum americicanum Araucaria biddwwillii Zanthoxylxylum simulans Arauuccaarria anganguusstifoollia uta graveolensens R Larix ggmmelinii a quasassioides Picrasm Larix decicidua s altissima Ailanthu Larriix kae ensis mpmpfeferi Toona sin LLariarix na laariricina Rhus typphihi ScSc i hihisandsandrara cchhi us potpotananininii inensnensisis RRhush Smimilalax hishisp rommaatica pidaida Rhus a AArrististolochc ntisccuus hiiaa mmaacrcro isttacia le ArArististolo phyllaylla PPis tale chia tommen ron oorriientaleen Chhimim entosa icodend ttaa oonannanth Tooxicodendx ia coorrddaa CCala us praaece Tiliail s lyycaanntth cooxx phhylloyllo CCa us fllooridi iliailia platypplaty alylyccantn duuss T ricaicana thus c iaia amamere Peumumu hinenshinensis TiTill lobloba uss bolbol is wia bibi CCiinnnn dus Grreew lba amamoom tutus a LaLauru umum cama ucalypyp us ruus non mpphoho E lobulul Pe obiillis ra ptutus g s rsseeaa ucacaly itritriinnu Sa boborbo E on c ssafaf ninia lisistetemom ananicica LiL rrasas ala CaCal geelll indeer lbibidumu iaia magma nanatata ra ben m ucuchshs pin Asim zzoo F ylealea p ica inana in taptaphyh olch Liri trrililoobba S ea colchicac a ododendren a phyl maldald Liri droo StaStaphylea buuma a riodiode n ttuulil yleaea rifofoliali MMa ndrdro ipipiferfe taphyph ea ttr aggnon on cch ra StaS hylel cox MaM olilia hinin Sttaap raaecoe aggno obob ensnse uuss prp siiss nol ovavatta yur nenns Mag ia tritr a tacchyurh chchiine lia nonolia ippeta S rus efolo MagnolMa lia m la hhyyuu nquefnqu ta gn ac taacc quiqui ser M olia ropoph S sus s in s aggnon sstteel hyyllal ciss susu nssii MMag oliiaa a llata a enoco ocicis rree ag ac rthenh eenn amu erra MMa nooll cuum PaPar arth itis a ininiffe agng ia dde iinatana P ViV tistis v attaa M nooliali ennuud ta Vi pidid agag a k daatt cus llala PPa no obub a ttrrici phy acc lliiaa s us us loophyl a hy sal isisss egalega ian Dece ssaan liciiciffo nnooc m gginianain is caia nd oliaia thhe sis m viirr nal AkebAk isnsnea ra t aarrt lopsop lis er a eb ea eerrmim P ele me is vernav ic SinofS iiaa insiinsi inna Amp maamelisel ino quin gngni alliiss Haam amem japonponi lllliiss MenM frraan a is am is mo en nc ttaa H el liiss ica C ispep hehe am el rrssica le err ttiiaa amamelam amem pepe sis CCl mamat m chinch H mma ia n le tis umum in Ha rotot innene N m liig canc en arrr s iana Naand aatti gu anad sissis P sis ti r nd s t stict ad opps onont ajoor Be in eernr icifif enen yc em maj i rb a d niffl ololi sese S ququem la m ryyi Be e doom lorra a aacc gil enren r ris t m a s j h ta B beer th es isi hergiher niaa iccaat er risis a unb titic iiopsops Fot soni sp sisis B ber a e a ot l iiss n erberirb iiss quiu rrgg aarrotrr owilsoowi nensne rraa BBe eri mmic iffolo ii P in ops si flo e i liumi S rryloyl isis ciif m B rb s bu cropro um o psps ucu u erberie er bu p CoC lo pa icuic is rb is x hyly ryl is on allis NNyss e jaeja iffoo lla o s p t a yss ris e lliia CCory op ja ien lua D v sc ylol m rien ifl a a s u hk orry llumu r oro acifac sa JamJ vididi syly lgag eae C ylly aar yr o am ia lv arir an ph b st ttiicco la DDeu ini aatti is a di ambm r st u ulau e eess nvvo icca ci iidd ar ffrru sc m D u ia o a er u bba u a u eeutztz ama lulu C iiquq m ssu mma rur H u iiaa m cr L ida iiaa ia ub uumm y tzia scas e aat ui on nni r in HHy d ia ca riic ta iqiqu e o ess p um rraa ddi b cana L a eo ibeb allp u yddr n iscolos rara n Phylogenies of 1369 temperate species PaP PaePa R s a nth m H ran g c a Fig. 2 an e a u y e o ib ccanthlatumat H dr geg a lo R ia uul um y a e mam r did ic m H d ng a a s c eos u y rangera eea ppani crcr as tet grru m HHy ddra n a a o iibebe fasf ac i u y ra gea q ni ph R s r nig e m HyH drd n a u cu hy brb s n ur y rana g e l llall ibesibe s be aaureu tum CornC d n e ara rc aatata a R iibes catu ra geg a rbo iifof ibebes R s ica tumtum C o e a o RRi bbese a rn n a p s rree liaia i sp icic um C o us ge p p s R s sp r eume re C rn a e eerr c e vvaa in C orno u k a ttii a een (a) or 501 deciduous-only species (b) with ib i lgalgar m C rnus s ouo a olo ns d n l s RiR d guin u liiumu CCor o u n u o a s n v ta rn s u sa m ri e angua ifolo otao or u flflo t s ibibe s m s ib ui CCo n s m or ttaa a R s rurum us o CCo o usu r llliilii la t tu m ihho se rn s m id ibesibe s btb u u o u a a u o r q n ttaa C rrn of s R igg stst q e o nus seris f LiL m in llaa is C r us a ficci um u m sis u s i oor n lb i trtru iguig ru c nnsi i C r u s nnal s L t m ti wiw Cor o nusn s m eri a al s u eeti re o a C rnr u i igu u r r u r sas Co or n s ma c s LLigus iggustrumststru a ara lo s C u w a eea LiL u gga m m illl riris o nusn ams c a autumn colours of leaves indicated by colours. n a vi a tata CoCor r u a ror ig i o v ga C n s s l o L r a a kko a llg a C u a terte p y g u m D rnun amo ngun h SSy in a g v blbla umu o s r g ri yly ringr gga inng o h ea S rrnn us r u l y in ri a c a iospyrio s a o llaa rinr yry g a tiich aaea s u cec m in S y S g p m a StS ttee s a e e n nnga oistis oop im p sps um S ring ri d r s s S tee w y f p mom a yri ri u ns ara r o e isissi e s A t w oos eemi o SyS Sy m d eeu d d e rt s lo m ririf s ridi pe iid ns AcA cct wwar aar tia l f a llu ia ir sspensao a A t r i ol vi u eo c ina Ac cti in art titi otus n k y ththia v rre iican a Acti ti a p tus a iiaa s h y y t utina c A ctinidi nidn idid ti t phylp ia a llyll u lut Sty t a m e rsyrs h iia s i r ni na a St c ini id ia ro Pagel’s = 0.6 for both phylogenies. ioo t h frutf a t p o li y t h vevel ana n P in iai e p va c a C y n d a c s n p Fo rsys ph m s l i iana dad C tet rrax idi ia k h eudoe aad e belb sysyt u y r iiaa e a d a k or r ia u erie n n l r E l in u d tit A nu ssy i e rostr x j iaia ararg o iolo FoF o s iin in n l u olo P nkink o lol e d e F e m o funf if ioior C t s jap o oc l la m x nnn r t s ia Ch i h a mme g nsisn p nen aaxinus ama ro lls L ere rra t m s c hha ta a r a L ia a y p e uut ama tat as pe s carc prop la e folia us a V h ri r o t iki is a FFr pe c i Va e a i ntn a a l a kt m n J s u s s ttifol nus nan s V s a nnic aan t n s xce s r V a u ma j t x nandr a ele o u iinu u a si Va j h rrb ic usu ex u oorn i si ta V c a b a l F nun x e E a c uus h uus l i in inusinu in g s ldid n ata E c o pop oor nd ia a x s en la us a ccc e s iis s x raxr xix x ngun u olo R ini t o r spi u ul c us s m c c n h d camc e p a F a a a n b inne i R i nin r ra r r iinusn a es a R h cicin n iui o a am a i a ebe h Rh p u i d FFr FFrax FraxinuFra x s i us s h ood i e ph c Phylogenetic data from Zanne et al. (2014). FFr ded a i RRh e n u m u ph us cchi aax u axi oos R ho odo d a r raxin s etusus oi i ei R trt iu m f p n s rangu irginicirgin i ii h o d r o inui s rret cic R R o u n F FFr d v vat ge s a hoh odo u m n a x uus nin R R dod o d c e adrang s e oovat ng a hho m l nnu n o e s iai i R o d o d e ig t a nunus i ppe l iii R h odod oryo g a c u rra i ua u un geg d odo m c x us a a RhR hho ded een n n ini s r to o d odo d n r n n a l F q i ri RhR hho o d ata aaxi hush thu gn fof ssa Rh o e ndr d yr ymb e lpa n i v a R h d i o lycl r t a s far a o od d ene g m yc t a b o e R ho d ndn d r g ssia u bi alpa fa eentosa avida i R o odeno tti s F us qu p m a R Rh ho d e nd o r Frax anthan l pa bu rni b Rh od ooden dr r i i s a nnieri oli a R h d d u lll b u a u ant m d i u I Rh d een ndronn o n l n n ern d o f t s ho o ood dr ron a i Bac Il do d e u o m nan la n alp l r a ho oodo odeno n o n m l pa talt o d A B l do d d r y a t lt a a e o d ye s Cat cifoli a El A e ho o e s s a j glog a i E n o a ic d a E o d d d n q t io alpa to o aecaeca eend r axinus a a l n desd ida E a ddod o q e u a hio al t ra x o dde nd drod o a eje b r inati a SSa r o r n m rraxi t ma V ddo e ro hionanh l bbod zi n CaC a r a VViburn le cch d m u d m CatalC a ia p pip im s aal per ddod o d o uin gr V l eut e ro n vi C j iipli g oi Vib F a V eeu o dden n ja i c WWei a p d c WeWeigela hort V d l o e a m s sis l a ib c e in C r D nnia t i at d r n o

i a es a ib l o n u eje i lla t u uc CCatalp WWeigela u t ta a i e i W ibu i d W W d d

n W u erny on l a m v CatCa d a m s a iburi h ndn rron n e Leaf colouration data from own observations. b i i a el le nnata e b d end dr t lmoil ni p yl n a d quq w h y i t s urnu bar w i a e b n r o o c s r ipe s enendr i d i s u t e n uddl rprpa cac g titis h e n cu his ki h a l k ei her d i s m urnu s l m d r i h n o u a o th e u b h e r r u o u o e ny deg e e o e h ena e n d u rn ow i e u um l e d ccic ic e um l ica rn c u ses a o n c r o n c on acu i ri r o c o ulea i e r r c BuB a a m u s dddleja alte mim cra u rnum lo u ige e nnd d n c e n r l ig r u e ph g nt i i c p g i i gelg viv c ra i o r e r la is d rro o m e u uddl c d o i g o nu g g n a era nig a n g nnum ro ata s faf mo b u m u uc n um aalo i n i n u ia n n r i dr r o f a o c g i iu a a i ap a a i umu cu llica eera rra i acila o lul e n au a ip eer oc r forf d o e o s ia cerc j l o u ll t f e h e e BudB a e o l a e u t steumst a l r r e m BBu sk e o l u n o tti j lp i u r oon ron c s m h ssm u o r o l a cam i c u la l a a een u lal l P i la m n a l oon v o a c l l a co i acu l m c i o gr n r a ycyciu ereripl ni yys m n h mirnow e a a s a d ax ic a a a r p u a ph ag m ooc m g c n m h n m m l m me m rre n r o n a n ra ta m i ovskiaov at rrag m i tut CalC L a lo o o ffor m n e m P l ei hytu max mo a ac ho c u o t a a r o n yp t ppra ca n imi sts o i i oon inv f ca cccu i h m h d j m ere m h a a tat c

s p a f ly pru a ra a p rn o an omm LoL r l p m o a xylxylo a le i m inv ra u l ccus v yt m r c e r ra n e igigr c t L c c o ra r t o E a i e a l er o r i era a u eer r s n p u t p a m icly tunei p a a ic e e e i rar e e rt ic nnt u rrles u ifoi ig o um PPer r ig u

o r r c cec ra r r x lll ana uc cce lul ra fol u nic a s r y a o l y i ra i e le m z ryt c r i ce a r e c m t u s se wi m ice eensis nif es a d e n o n e r ici i e e i e t y n m i on a s i llo E mowim n c ni n i c ll cec n e s lia l o n ns s e c a ico cer r i ii i cer e n s r i l thr u n p s i LLo a g c ice o o i ici ce o o on cera i a h o m a i a e c a oolium ii i ca w o i c i cer o m n oni o e l m oonicera ca ono eebol os n iic o su n x s LoL n x ru a LLo a u n k is li LLonicera alpi a For (a), data from Archetti (2009) were added w o t LLonicer p i L e t b L l su u o n rar um oon u c r r s L on d m b ol um m p o m LonLo LoniceraL on e i L m hhu

i a c LLoni s m u o u

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i c m e i e i n r

i aan a nne t o o n ens p L h u n e p s si

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y to include evergreen species. S absorbed by Chlb, thereby protecting leaves from photoinhibition the efficiency of nutrient retrieval from senescing autumn leaves (Gould et al., 1995). Experiments in a North American dogwood (Feild et al., 2001). Over the past two decades > 50 studies have species further supported the hypothesis that optical masking of explicitly tested for photoprotection in a wide variety of systems to Chl by anthocyanins could reduce the risk of photo-oxidative explain seasonal, developmental, and inter- and intraspecific damage to leaf cells as they senesce, which otherwise might lower differences in leaf pigmentation (Gould et al., 2018, table 1). In

Ó 2019 The Authors New Phytologist (2019) 224: 1464–1471 New Phytologist Ó 2019 New Phytologist Trust www.newphytologist.com New 1468 Review Tansley review Phytologist

1.00 c. 30% of these studies, however, the physiological responses of red and green leaves to light were comparable or else, the anthocyanin- red foliage appeared to perform worse than green foliage. The explanation for this probably lies in the context-dependence of 0.75 anthocyanin benefits. In order to test for such context-dependence, Gould et al. (2018) compared the stress responses of WT (Col-0) Arabidopsis thaliana, Leaf colour which had only trace amounts of foliar anthocyanins, with stress 0.50 Yellow responses in the anthocyanin-rich pap1-D mutant, and, in some experiments, the anthocyanin-deficient ttg1-1. They recorded Red Percentage effects of light quality, light intensity, air temperature and duration of exposure on quantum yield of photosystem II (ΦPSII) in leaves 0.25 47% of different ages and compared pigment profiles before and after treatment. Although similar in anatomy, Chl content and xantho- 38% phyll cycle activity, the anthocyanin-rich pap1-D mutants were less 20% photoprotected than WT Col-0 (with only trace amounts of 0 anthocyanin), but only when given at least 2 h exposure to North America Asia Europe saturating light at 10°C. Differences between the responses of red Fig. 3 Percentages of yellow and red species per continent for 711 species and green leaves were greatest in older plants that were given monitored in the Munich Botanical Garden. North America, Asia and Europe protracted exposures to high fluxes of cool-white light (5600 K) at are significantly different in their occurrence of anthocyanin (chi-squared chilling temperatures. This shows that anthocyanins’ photopro- < test: P 0.001). tective function is greater at lower temperatures and under higher light intensities (Gould et al., 2018, figs 2–4). Based on these findings, Gould et al. (2018) suggested that photoprotection by anthocyanins should be of greatest adaptive value for plants receiving prolonged exposure to intense cool-white light or natural sunlight at low temperatures, when the capacities of other photoprotective mechanisms are insufficient. Anthocyanins may, in addition, serve as useful sinks for excess photosynthates produced under saturating light; the close association between anthocyanin accumulation and sucrose content suggests that this could be their other important role (Gould et al., 2018).

IV. The adaptive value of colour-changing leaves Two main adaptive explanations for red or yellow autumn colouration have been put forward: the coevolution hypothesis and the photoprotection hypothesis. According to the coevolution hypothesis, bright autumn colours are warning signals to insects, especially aphids, that lay their eggs on the trees in that season. ‘If the colour is linked to the level of defensive commitment of the tree and the insects learn to avoid bright colours, this may lead to a coevolutionary process in which bright trees reduce their parasite load and choosy insects locate the most profitable hosts for the winter’ (Archetti & Brown, 2004, p. 1219). This explanation was first proposed by William Hamilton (Archetti, 2000; Hamilton & Brown, 2001; Archetti & Brown, 2004; Archetti et al., 2009). The two fundamental predictions of Hamilton’s signalling hypothesis are that (1) trees use leaf colouration as an honest signal to communicate their defensive capacities to insects and (2) differences in leaf colours change the behaviour of insects. Over the Fig. 4 Colour change in Acer rubrum and Liriodendron tulipifera individuals past 18 yr, neither of these predictions has been supported by monitored in St Louis, MO, USA, 2007–2018. The timing of colour change in both species is highly consistent among years, with the intensity of the experimental or comparative data. On the contrary, there is anthocyanin colour in the maple trees depending on the preceding night evidence that the defensive commitment of a tree or a tree branch is frost; no such intensity difference could be seen in the carotenoid colour of negatively correlated with anthocyanin production. This is because the tulip tree. Photo: P. Renner, 11 November 2013. phenylalanine precursors used in anthocyanin synthesis cannot

New Phytologist (2019) 224: 1464–1471 Ó 2019 The Authors www.newphytologist.com New Phytologist Ó 2019 New Phytologist Trust New Phytologist Tansley review Review 1469 instead be used to produce other flavonoids of higher defence value Arabidopsis comes in. Their data show that anthocyanins’ (e.g. tannins, lignin, flavonols or flavones) or in the synthesis of photoprotective function is greatest at lower temperatures and hydroxycinnamic acids esterified to cell walls for reinforcing under higher light intensities when the need to quench excess light mechanical barriers against herbivores (see Meyer et al., 2006, for exceeds the cells’ capacity for thermal energy dissipation (further the broader context of these trade-offs). Indeed, a study on Norway relevant experiments are cited in Section III). One would therefore maple (Acer platanoides) found that anthocyanin production was expect that red and yellow autumn leaves should be most common highest in trees with the lowest defensive commitment, such as in species exposed to sudden cold spells and high light intensities. partially dead trees with low leaf N content (Sinkkonen, 2008). This expectation can be tested against climate and solar Autumn pigmentation is thus not an honest signal of defence irradiation high-resolution data on the long-term monthly commitment. averages of temperatures and incoming radiation in North The largest study designed to test the second prediction (that America, Europe and Asia, the regions that are most relevant to colour affects herbivore behaviour) used Betula pendula genotypes the question of the adaptive value of autumn colouration in with green or yellow autumn colouration and Euceraphis betulae, arborescent species. The temperate deciduous forests of North one of the most widespread tree aphids in Western Eurasia America and Asia receive higher solar irradiation during autumn (Sinkkonen et al., 2012). The study failed to find a significant than does Europe (Fig. 5). In North American deciduous relationship between genotypic variation in leaf colour reflectance forests, the average daily short-wave radiation in September is or leaf attractiveness and the numbers of nymphs, males, or winged 168 W m 2. In Asian forests, it is 165 W m 2. By contrast, with females on the respective trees. It appears that autumn-migrating only 114 W m 2, European forests receive 32% less irradiation aphids do not automatically concentrate on trees with the most during autumn than North America (Fig. 5b). Also, for shrub-/ attractive (yellow) leaves, but also use other cues for their grasslands, autumn irradiation is highest in North America orientation, such as female pheromones in the case of male aphids. (190 W m 2) and lowest in Europe (141 W m 2), with Asia In a permutation of Hamilton’s hypothesis, Lev-Yadun & intermediate (162 W m 2; Fig. 5c). Holopainen (2009) proposed that the solution to the problem of Intercontinental differences in both leaf-out and Chl breakdown the origin of red autumn leaves in general, and their limited times between the arborescent species of ENA, Asia and Europe distribution in Northern Europe in particular, is the difference in have been found in studies that compared species from the three the extinction histories of trees in ENA and East Asia (lower regions grown under identical conditions (Zohner & Renner, extinction rate) and those in Northern Europe (higher extinction 2017; Zohner et al., 2017). Experiments on 396 species from East rate). ‘Not only did trees with red leaves mainly become extinct in Asia, Europe and ENA revealed that the length of the vegetation Europe, but when many of their herbivores became extinct, the period, which is the time between leaf-out and leaf senescence driving selective agents for red autumn colouration also declined. (measured as 50% colour change and/or leaf abscission), of ENA [...] Thus, the anti-herbivory component of the character of red species is 3 wk shorter than the vegetation periods of their European autumn leaves partly reflects anachronistic adaptations to past and East Asian relatives, because their leaves flushed 9 4 and faunas, many of which became extinct’ (Lev-Yadun & Holopainen, 13 4 d later and senesced 9 4 and 11 4 d earlier (Zohner & 2009, p. 510). This strikes us as circular-sounding or, to paraphrase Renner, 2017). The authors attributed these regional differences to Olson & Arroyo-Santos (2015, p. 167), ‘this trait was shaped by higher interannual temperature fluctuations in North America selection in unseen ancestral populations and this selection must compared to Asia and Europe (see fig. 1a in Zohner et al., 2017). have occurred because the trait is present’. Less predictable springs and autumns apparently have favoured Under the photoprotection hypothesis, the yellow carotenoids more conservative (safer) phenological strategies in North America and the red anthocyanins both function in photoprotection compared to Asia and Europe. (Section III), which is beneficial because it helps extend the period Given these regional differences in the leaf-out and Chl of nutrient resorption, especially N and phosphorus (Feild et al., breakdown strategies of Northern Hemisphere trees and shrubs, 2001; Hoch et al., 2001). Resorption from senescing leaves requires we believe that the higher frequency of red leaves in North America synthesizing numerous enzymes and regulatory elements involved compared to Europe reflects the combined effects of higher average in the degradation and remobilization of leaf nutrients (Hoch et al., autumn irradiation and earlier initiation of senescence when 2001; Landi et al., 2015). Reflecting this, leaf respiration continues intensities of solar irradiation are higher (Fig. 5). Additionally, unchanged or at an increased rate during senescence (Hoch et al., North American species likely experience more cold days during 2001) because as chloroplasts are getting impaired, the remaining the senescence process than do Asian and European species because organelles acquire novel functions, particularly the mitochondria of greater temperature fluctuations (above). Because the environ- (Keskitalo et al., 2005). In essence, senescing leaves need to retain, mental conditions under which the photoprotective benefit of leaf at least in part, their defence potential against both ROS and toxic pigments is greatest are precisely conditions of low temperatures lipid peroxidation products to successfully dismantle their photo- and higher light intensities (Gould et al., 2018), the higher synthetic apparatus during senescence in a relatively safe manner. frequency of colour-changing species in North America and Asia Although the photoprotection hypothesis has solid experimental (and their rarity in Europe) exactly match theoretical expectations. support (Section III) it fails to explain why red and yellow leaves In addition, the on-average 3-wk shorter vegetation periods of ENA should be more common in ENA and Asia than in Europe (Fig. 3). species (Zohner & Renner, 2017) may place an additional burden This is where Gould et al.’s (2018) experimental evidence from on American species of extending the period during which their

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(a) 300 250 200 150 100 Boston Rome Shenyang 50

) 0

–2 (b) Deciduous North AmericaEurope Asia 200 forests

150

100

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September SW radiation (W m (c) Temperate 200 shrubland

150

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0 –100 0 100 Longitude Fig. 5 Longitudinal differences in autumn solar irradiation. (a) Average (from 1901 to 2010) daily short-wave(SW) irradiation for the month of September. Data on SW radiation from Beer et al. (2014). (b, c) Continental differences in autumn SW radiation for deciduous forests (b) and temperate shrub-/grasslands (c). Error bars refer to 95% confidence intervals. Information on spatial distribution of biome types from Olson et al. (2001). photosynthetic tissues are protected from ROS by anthocyanins so anthocyanin should be less beneficial in nutrient-rich sites. This that nutrients can be resorbed. theoretically expected trade-off between investment in anthocyanin production and benefits from recycled nutrients receives some V. Outlook support from the observation that plants living in symbiosis with N-fixing bacteria, such as alders (Alnus), do not produce autumn This review sets up a new framework for an evolutionary colours (Archetti, 2009). A global test of this hypothesis is still understanding of autumn colouration. To test whether our missing. photoprotection-cum-regional-climate hypothesis explains spa- tial patterns of autumn colouration, species-level colouration Acknowledgements data need to be linked to species distribution data. Databases are available (e.g. GBIF for species occurrence points or GFBI We are grateful to Luc Bidel, Christian Jay-Allemand and the for abundance-weighted plot-level data) that would allow such reviewer Marco Landi for their plant-physiological expertise, which analyses. Also, a global map of solar insolation for the relevant greatly improved our manuscript; to the ecologist Simcha Lev- autumnal period is needed. The relevant period needs to be Yadun for his review; to the Editor Marc Rausher for his comments; defined based on regional leaf-senescence times obtained from to Emma-Irene Arndt, Anya Goncalves, Andrea Lang and Anna remote sensing data. Having data on the relevant autumn Satzger for gathering information on colour change in the Munich period will also allow testing for the effects of cold days Botanical Garden (autumn 2017); and to the DFG for financial during leaf senescence. support (RE 603/25-1). Another possibility to test the photoprotection-cum-regional- climate hypothesis would be to compare nutrient resorption ORCID strategies. In species (or populations) strongly limited by N, one would expect higher selection for efficient nitrogen resorption in Susanne S. Renner https://orcid.org/0000-0003-3704-0703 autumn than in species less limited by N and thus higher Constantin M. Zohner https://orcid.org/0000-0002-8302- investment in anthocyanins. Conversely, the active production of 4854

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