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1 Soils&and&nutrient&cycling&in& tropical&forests& Carlos&A.&Sierra& Max&Planck&Ins;tute&for& Biogeochemistry&

2 Outline& Nutrient&plant&demands& The&biogeochemical&heterogeneity&of&the& tropical&biome& Humans&and&the&cycling&of&nutrients&

3 Essen;al&elements&for&life&

4 Essen%al(Nutrients(of(Plants( & ( ( (Chemical ( (Atomic(Ionic(forms ( ( ( ( (Approximate(dry((( (((((Element (((( ( (symbol( (weight(absorbed(by(plants ( (concentra%on (((( Macronutrients&&&& Nitrogen & & & &N & & &14.01& & &NO 3I,&&NH4 + & & & & & & &4.0&%& Phosphorus & & &P & & &30.98& & &PO 3I 4,&HPO 2I 4,&H 2 PO 4I & & &&&&&&&0.5&%& Potassium& & &&& &K & & &39.10& & &K + & & & & & & & &4.0&%& Magnesium & & &Mg & &24.32& & &Mg 2+ & & & & & & &0.5&%& Sulfur & & & & &S & & &32.07& & &SO 2I 4 & & & & & & &0.5&%& Calcium & & & &Ca& &&&&&&40.08 & &&&&&&Ca 2+ & & & & & & &1.0&%& Micronutrients, Iron & & & & &Fe& & &55.85& & &Fe 2+,&Fe 3+ & & & & & &&200&ppm& Manganese & & &Mn & &54.94& & &Mn 2+ & & & & & &&200&ppm& Zinc & & & & &Zn& & &65.38& & &Zn 2+ & & & & & &&&&30&ppm& Copper & & & &Cu& &&&&&&63.54 & & &Cu + 2 & & & & & &&&&10&ppm& Boron & & & & &B & & &10.82& & &BO 2I 3,&B 4 O 2I 7 & & & & &&&&&&&&&&60&ppm&& Molybdenum & & &Mo & &95.95& & &MoO 2I 4 & & & & & &&&&&&2&ppm& Chlorine & & & &Cl & & &35.46& & &Cl I & & & & & &&&&&3000&ppm& Essen.al, Carbon & & & &C & & &12.01& & &CO 2 & & & & & &&40&%& Hydrogen & & & &H & & &&&1.01& & &H 2 O & & & & & &&&&6&%& Oxygen & & & &O & & &16.00& & &O 2,&H 2 O & & & & & &&40&%& & &Plant&;ssues&also&contain&other&elements&(Na,&Se,&Co,&Si,&Rb,&Sr,&F,&I)&which&are&not&needed&for& the&normal&growth&and&development.&

5 Primary&func;ons&of&elements& Func%on( Elements( Chemical(form( Examples( Structural& Electrochemical& Cataly;c&(acid& base)& Cataly;c&(redox)& H,&O,&C,&N,&P,&S,&Si,& B,&F,&Ca& H,&Na,&K,&Cl,& HPO 4 I2 & Zn,&Ni,&Fe,&Mn& Fe,&Cu,&Mn,&Mo,& Se& Organic& compounds& Free&ions& Complexed&with& enzymes& Complexed&with& enzymes& Biological& molecules,&cell& walls,&;ssue,& skeletons,&etc.& Message& transmission,& cellular&signaling,& energy& metabolisms& Diges;on,& hydrolisis,&po 4 & removal&in&acid& media& Reac;ons&with&O 2, & N&fixa;on,& reduc;on&of& nucleo;des& Sterner&&&Elser,&2002.&

6 Chapin&et&al.,&2011&

7 Nutrient&supply&constraints&the&produc;vity&of& terrestrial&ecosystems& Chapin&et&al.&2002.&

8 N,&P,&and&K&are&the&most&important&nutrients&that& contribute&to&plant&growth&in&agroiecosystems& Source:&FAO&2005&

9 Justus&von&Liebig's&Law(of(the( Minimum& Plant&growth&is& controlled&not&by&the& total&amount&of& resources,&but&by&the& scarcest&resource & y =min{f N (N),f P (P ),f K (K),...,f L (L)}

10 Nutrient&supply&constraints&the&produc;vity&of& ecosystems&

11 The&supply&of,&and&demand&for,&N&and&P&are& usually&in&close&balance& Davidson&&&Howarth&2007.&&

12 N&and&P&are&the& most&important& nutrients&for& biological∾vity& Sterner&&&Elser&2002&

13 Different&biological&molecules&have&fixed&stoichiometric& propor;ons& Sterner&&&Elser&2002&

14 C,&N,&and&P&can&be&found&in&constant&propor;ons& in&the&oceans&

15 N:P&ra;os&might&be&indicators&of&nutrient&limita;on& in&terrestrial&ecosystems& N:P&<&14& N:P&>&16&

16 N:P&ra;os&appear&to&indicate&P& limita;on&in&tropical&ecosystems& Hedin&2004&

17 Soil&nutrients&change&with&soil& development& Lambers&et&al.&2008&

18 Soil&pH&and&nutrient&concentra;ons&

19 Phosphorus&in&soils&decreases&with& ;me& Walker&&&Syers&1976&

20 Test&of&the&Walker&&&Syers &model&in&a&longiterm& chronosequence&in&hawaii& Crews&et&al.&1995&

21 Test&of&the&Walker&&& Syers &model&in&a&longi term&chronosequence& in&hawaii& Crews&et&al.&1995&

22 Changes&in&nutrient&limita;on&with&soil&development& Vitousek&&&Farrington&1997&

23 Take&home&messages& Nitrogen&and&phosphorus&are&the&main& nutrients&that&control&plant&growth& N&and&P&limits&plant&produc;on&in&a&synergis;c& fashion.&addi;ons&of&n&must&be&balanced&by& more&p&and&vice&versa.& P&limita;on&is&related&to&soil&development.&As& a&consequence&tropical&forests&are&believe&to& be&p&limited.&&

24 The&biogeochemical&heterogeneity&of& the&tropical&biome&

25 Source:&hmp:// soil4505/doc/unit10af.htm& Soils&of&Africa&

26 USDA&soil&orders&and&soil&development&

27 Soils&of&the&Amazon&basin& Quesada&et&al.&2011&

28 Soil&development&in&the&Amazon&

29 Oxisols&(USDA)&or&Ferralsols&(WRB)& Highly&weathered& High&clay&content&but&of&low& ac;vity&(1:1itype)& Deep&soils&>&20m& Good&drainage& Low&natural&fer;lity& High&concentra;ons&of&Fe& and&al&oxides& Moderately&acid&

30 Ul;sols&(overlaps&with&Plinthosols)& Highly&weathered& Clay&weathering&and&transloca;on& Acid&B&horizon&with&<&35%&CEC& No&calcareous&material& &<&10%&weatherable&minerals&in&topsoil&

31 Spodosolos&(Podzols)& Intensive&acid&leaching& Subsurface&horizon&with&illuvia;on&of& organic&mamer& WhiteIsand&horizon&of&eluvia;on& Some&of&the&most&nutrient&poor&forest& in&the&tropics& do&nascimento&et&al.&(2004)&

32

33 But&tropical&forests&are&supposed&to&be&the&most& produc;ve&ecosystems&on&earth!& Beer&et&al.&2010&

34 Tropical&forest&paradox& The%tropical%rainforest%thus%has%a% rela2vely%rich%nutrient%economy% perched%on%a%nutrient7poor% substrate % Whimaker&1975&

35 Primary&produc;vity&is&controlled&by&the& availability&and&the&efficiency&of&resources& NPP =min{l LUE, N NUE,W WUE} L = Light N = Nutrients W = Water Agren&&&Andersson&2012&

36 Strategies&to&increase&NUE&in&tropical& forests& Retransloca;on& NIfixa;on&and&symbio;c&associa;ons& Root&mats&

37 Limi;ng&nutrients&are&retranslocated&before&leaf& abscission&& Fife&et&al.&2003&

38 Roots&do&not&appear&to&retranslocate&nutrients& Gordon&&&Jackson&2000&

39 N&fixa;on&in&tropical&forests& Hedin&et&al.&2009&

40 NIfixing&organisms&

41 FreeIliving&N&fixers&in&a&tropical&forest& Reed&et&al.&2008&

42 Legumes&are&important&NIfixers,&but&not&always,& not&everywhere& Barron&et&al.&2010&

43 Roots&release&organic&substances&& (chelates&and&phosphatases)&to& mineralize&p&

44 RootImats& Jordan&&&Escalante&1980&

45 Strategies&to&increase&NUE&in&tropical& forests& Retransloca;on& NIfixa;on&and&symbio;c&associa;ons& Root&mats& LongIlived&;ssue&(leaves)& Defense&compounds&against&herbivory& Adjust&shoot&to&root&ra;os&

46 The&biogeochemical&heterogeneity&of& the&tropical&biome&

47 Wood&produc;vity&seems&related&to&soil&age&in&the&Amazon& basin& Malhi&et&al.&2004&

48 Soil&P&may&explain&differences&in&aboveground&wood& produc;on&

49 But&aboveground&NPP&may&not&tell&us&the&whole& story& Aboveground NPP Belowground NPP Productivity (Mg C ha 1 yr 1 ) Productivity (Mg C ha 1 yr 1 ) Clay soils Sandy soils Clay soils Sandy soils Total NPP LAI index Productivity (Mg C ha 1 yr 1 ) LAI index (m 2 m 2 ) Clay soils Sandy soils Clay soils Sandy soils Jimenez&et&al.&2014&

50 TakeIhome&message& The&tropical&biome&is&highly&diverse&in&its& biogeochemistry,&but&most&regions&have&highly& weathered&soils& Highly&weathered&soils&suggest&P&limita;on&in& these&forests,&however&they&are&s;ll&highly& produc;ve& This&paradox&can&be&resolved&in&terms&of&the& efficiency&of&nutrient&use&

51

52 Humans&and&the&cycling&of&nutrients&

53 Shiuing&cul;va;on&

54 Shiuing&cul;va;on&

55 Uhl&&&Jordan&1984& Ecosystem&recovery&auer&slashIandIburn&

56 Ecosystem&recovery&auer&slashIandIburn& Uhl&&&Jordan&1984&

57 Recovery&cycle&and&soil&proper;es& Chazdon&2014&

58 Biomass&recovery&in&tropical&America& Poorter&et&al.&2016&Nature&530:&211I214&

59 Biomass&recovery&in&tropical&America& Poorter&et&al.&2016&Nature&530:&211I214&

60 What&are&the&effects&of&large&scale& deforesta;on&on&nutrient&cycling?&

61 Effects&on&aqua;c&ecosystems&

62 Important&amounts&of&N&are&removed&by& deforesta;on,&but&it&can&recover&in&secondary&forests& Markewitz&et&al.&2004&

63 P&is&also&removed&but&an&important&por;on&is& retained&by&minerals&in&the&soil& Markewitz&et&al.&2004&

64 N&recovers&auer&agricultural& abandonment& Davidson&et&al.&2007&

65 Davidson&et&al.&2007& A&new&paradigm&

66 N&fixa;on&seems&to&be&key&for& phosphatase&produc;on& Houlton&et&al.&2008&

67 C&stocks&can&recover&auer&decades&of& landiuse& TAGB (Mg C ha!1 ) TBB (Mg C ha!1 ) TDM (Mg C ha!1 ) Successional age (years) Successional age (years) Successional age (years) SOC (Mg C ha!1 ) TCS (Mg C ha!1 ) Proportion TAGB/TCS TAGB/TM Successional age (years) Successional age (years) Successional age (years) Sierra&et&al.&2012&

68

69 Con;nuous&cul;va;on&and&fer;liza;on&

70 However,&con;nuous&cul;va;on&is&not& sustainable& Stocking&2003&

71 Fertilier use (kg ha-1 arable land) Colombia China Germany USA World Year

72 Phosphorous&peak&

73

74 Phosphorus&use&efficiency&

75 TakeIhome&messages& SlashIandIburn&of&tropical&forests&make& nutrients&available&for&crops,&but&produc;vity& decline&over&;me& Regrowing&forest&can&recover&nutrient&stocks& over&;me& N&is&a&key&nutrient&for&forest&recovery& Current&fer;lizer&use&is&not&sustainable&

76 Topics&for&discussion& Environmental&consequences&of&largeIscale& conversion&of&tropical&forests&(oil&palm,&soy).& Sustainability&of&crop&produc;on&in&tropical& regions.& Possible&solu;ons&to&balance&food&produc;on& and&nutrient&loss.&

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