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Constituintes da MOS

LSN_5897 2-Constituintes Da MOS

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Constituintes

da MOS

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Ecossistema Natural Agrosistema

Restos deculturasLiteira

Resíduos org:agrícolaurbanoindustrial

Pluviolixiviados

Macro e meso faunae microrganismos

MetabólitosExsudatos

Raízes

Exsudatos

Raízes

CO2

Carvão

Conceitos e constituintes da MOS

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Acimasuperfície do solo

Abaixo

Vivo Morto

PLANTA

Exsudatos Raizes

ORIGEM DO MATERIAL ORGÂNICO

Pluvio-

lixiviados

• Liteira• Restos de

culturas• Resíduos

MACRO/MESOFAUNA EMICRORGANISMOS

Vivo Morto

Macrofauna

Mesofauna

BiomassaMicrobiana

Restos animais e

excreções

Restos de micro-organismos e

metabólitos

Conceitos e constituintes da MOS

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CONSTITUINTES

Conceitos e constituintes da MOS

Pluviolixiviados

Liteira, restos de culturas e resíduos orgânicos

Raízes e exsudatosOrganismos do solo

Substâncias não-húmicasSubstâncias húmicas

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CONSTITUINTES

Conceitos e constituintes da MOS

Pluviolixiviados

Liteira, restos de culturas e resíduos orgânicos

Raízes e exsudatosOrganismos do solo

Substâncias não-húmicasSubstâncias húmicas

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 Definição: Porção da chuva que chega ao solo passando pela copa da vegetação,pelas aberturas nas copas, gotejando das folhas, galhos e troncos.

Pluviolixiviados (throughfall)

Evapotranspiração

Interceptação

Armazenamento(nas palntas)

Infiltração

Absorção

Pluviolixiviados

Conceitos e constituintes da MOS

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Metodologia e exemplos

Coleta Preservação Análises

Tipos de coletoresColetor de chuva/pluvio lixiviados

Funil (16 cm )

Filtro de pol iester 

Garrafa coletora

Haste de metal (1,5 m)

1 m acima do solo

Solo

Conceitos e constituintes da MOS

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Tipos de coletores

Conceitos e constituintes da MOS

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Tipos de coletores

Conceitos e constituintes da MOS

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Exemplo de coleta de pluviolixiviados na Amazônia

Conceitos e constituintes da MOS

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Entrada de carbono

Floresta

0

200400

600

800

Jan Fev Mar Abr Mai Total

   C

  m  g  m  -   2

0

2000

4000

6000

   P

   l  u  v   i  o  s   i   d  a   d  e   (  m  m   )

PluviolixiviadoChuva

Conceitos e constituintes da MOS

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Fluxos no sistema florestal (mg C m-2)

Pluviolixiviado5277

1 m1 m

1 m

Escoamentosuperficial

173

Chuva3022

Lixiviado

7,34

Conceitos e constituintes da MOS

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2717,96860,53021,68920,8Total

278,4147,9346,3855,6Mai489,11109,5435,1189,9 Abr 

734,60206,3867,92243,8Mar 

409,77219,4510,04248,8Fev

806,07277,4862,23282,7Jan

mg m-2mmmg m-2mm

Entrada CChuvaEntrada CChuva

PastagemFloresta

Mês

Pluviosidade e entrada de C

Conceitos e constituintes da MOS

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CONSTITUINTES

Conceitos e constituintes da MOS

PluviolixiviadosLiteira, restos de culturas e resíduos orgânicos

Raízes e exsudatosOrganismos do solo

Substâncias não-húmicasSubstâncias húmicas

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Liteira, restos de culturas e resíduos orgânicos

 Ecossistema natural 

 Resíduos orgânicos

 Agrossistema

 Definição: resíduosvegetais e animaisdepositados nasuperfície do solo

Conceitos e constituintes da MOS

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Composição elementar e bioquímica

Água

75%

Matéria seca

25%

    A   g     u    a 

    7    5

    %

Celulose

45% Açúcar e gomas5 %

Lignina20 %

Hemicelulose20 %

Gorduras eceras 2 %

Proteína8 %

Tipos de compostos

Carbono

44 %

Oxigênio

40 %

Hidrogênio 8 %

Cinzas 8 %

Composição elementar

Conceitos e constituintes da MOS

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Tecidos vegetais

Carboidratos

Aminoácidos e proteínas

Lipídeos

Ácidos nucleicos

Lignina

Tecidos animais

Quitina (parede celular de fungos)

Ácido murâmico (parede celular de bactérias

Natureza bioquímica dos tecidos vegetais e animais

Monossacarídeos: glicose, frutoseOligossacarídeos: maltose, sacarosePolissacarídeos: celulose, amilose, hemicelulose e

ácidos poligalacturônicos{

Côr verde-azuladaindica presença delignina na epidermee córtex

Conceitos e constituintes da MOS

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Velocidade de decomposição dos constituintes orgânicos

Monossacarídeos

Proteínas

HemiceluloseCelulose

Lipídeos

Compostos fenólicos

Lignina

LentaLenta

RRáápidapida

Conceitos e constituintes da MOS

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Produção de liteira e restos de culturas

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                             P                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                        r                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                         o                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                    d                       

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                      u                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                ç                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                       ã                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                            o                        

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                            t                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                              o                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                    t                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                              a                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                l                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                            d                       

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                         e                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                     l                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                      i                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                            t                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                              e                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                              i                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                        r                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                         a                                   

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                           (                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                  t                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                  h                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                         a                                   

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                     -                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                1

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                         a                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                           n

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                         o                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                             -                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                1

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                            )                                                                                                                                                                                                                                                                                                    

0

4

6

12

20 40 60Latitude Norte/Sul (graus)

(Adaptado de Bray and Gorham, 1964)

EquatorialTemperado quente

Temperado frio

Ártico

 Produção anual de liteira de florestas em relação à latitude

Conceitos e constituintes da MOS

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 Dinâmica sazonal da produção de liteira

Final do “verão”(período seco)

Meio do “ inverno”(período de chuvas)

Final do “verão”(período seco)

Estações do ano

Acúmuloliteira

Conceitos e constituintes da MOS

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FlorestaLiteira

totalSomente

folhasFolhas

A. Primária (platô) 7,5 4,4 59B. Primária (várzea) 6,5 4,0 62

C. Secundária jovem 6,1 5,2 85

%t ha-1 ano-1

Matéria seca remanescente durante processo de decomposiçãoem relação ao peso inicial

Tempo (dias)

Estação seca Estação chuvosa

Conceitos e constituintes da MOS

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Quantidade de restos vegetais (matéria seca) de diferentes culturas

Clube Amigos da Terra, 1995

Soja 2.000Nabo forrageiro 4.000Aveia preta 5.000Cevada 2.000

Centeio 3.000Trigo 1.500Milho 6.000

Culturas Massakg ha-1

Palhada no campo

Conceitos e constituintes da MOS

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Composição dos restos de algumas culturas

Cultura N P K Ca Mg _________________________________________________ 

----------------%--------------------------

Trigo 2.80 0.36 2.26 0.61 0.58

Milho 2.97 0.30 2.39 0.41 0.16Amendoim 4.59 0.25 2.03 1.24 0.37Soja 5.55 0.34 2.41 0.88 0.37Batata 3.25 0.20 7.5 0.43 0.20

Beterraba 3.76 0.38 4.01 0.78 0.68Algodão 3.29 0.37 2.07 2.48 0.49Alfafa 4.63 0.48 2.76 2.38 0.66

 _______________________________________ 

Conceitos e constituintes da MOS

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Metodologias de quantificação e exemplos

 Ecossistema natural: liteira

Coletor de liteira em queda Coleta de liteira do solo

   2   0  c  m

Superfície

do solo

Estaca desuporte

80cm60cm

   1  m

7   0   c  m  

1  5    c  m  

Tecido robusto

Polietileno duro

Recipiente

plásticoremovível(material fino)

malha de nylon(1mm)

Tela de arameremovível

(material grosseiro)

Conceitos e constituintes da MOS

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 Agrossistema: restos culturais

 Exemplo: cana-de-açúcar

Corte sem queima x com queima Medida do volume de material coletado

Palhada ~ 14 Mg ha-1

matéria seca

Campos 2003

Conceitos e constituintes da MOS

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 Agrossistema: restos culturais

 Exemplo: plantio direto

Exemplos: Região Mg ha-1

Sul (Tibagi-PR) 10,5

Centro-Oeste (Rio Verde-GO) 5,9Siqueira-Neto, 2002

Conceitos e constituintes da MOS

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 Preparo das amostras

Moagem AnálisesSeparação por classes Secagem

Conceitos e constituintes da MOS

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Vinhaça

BiossólidoComposto de lixo urbano

Tipos de resíduos orgânicos

Conceitos e constituintes da MOS

C i i i d MOS

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Resíduos de cortume

Resíduos petroquímicos

Farinhas e resíduos frigoríficos

Torta de origem vegetal

Estercos

Tipos de resíduos orgânicos

Conceitos e constituintes da MOS

C it tit i t d MOS

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Composição percentual de resíduos frescos de animais

Fonte N P K Ca Mg

Gado leiteiro 0.53 0.35 0.41 0.28 0.11

Gado de corte 0.65 0.15 0.30 0.12 0.10

Cavalo 0.70 0.10 0.58 0.79 0.14

Galinha 1.50 0.77 0.89 0.30 0.88

Porco 0.58 0.15 0.42 0.57 0.08

Conceitos e constituintes da MOS

C it tit i t d MOS

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CONSTITUINTES

Conceitos e constituintes da MOS

PluviolixiviadosLiteira, restos de culturas e resíduos orgânicos

Raízes e exsudatosOrganismos do solo

Substâncias não-húmicasSubstâncias húmicas

C it tit i t d MOS

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Raízes

Conceitos e constituintes da MOS

C it tit i t d MOS

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 Produção de raízes pelas principais culturas tropicais

Prof. Raízes Restolho Total

(cm)

Trigo de inverno 0-30 0,69 0,45 1,14(não fertilizado)

Trigo de inverno 0-30 0,88 0,59 1,47(com N,P,K)

Aveia 0-30 1,00 0,57 1,57

Milho 0-80 1,04 --- ---(com N, P, K)

Soja 0-60 0,82 --- ---

Conceitos e constituintes da MOS

Conceitos e constituintes da MOS

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 Produção de raízes pelas principais culturas tropicais

Profundidade (cm)

0-4

4-9

9-1515-30

30-60

60-100

Total

Matéria seca t.ha-1

0,938

0,191

0,1400,150

0,175

0,147

1,741

Distribuição %

54

11

89

10

8

100

FUNDAÇÃO MS, 2000

 Brachiaria decumbens

Avaliado após 16 anos de formação e pastejo contínuo

Conceitos e constituintes da MOS

Conceitos e constituintes da MOS

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 Produção anual de biomassa de raízes com a liteira da floresta Amazônica

Biomassa aérea 7,6 121 2

Biomassa de raízes 8,0 190 9

Mg.ha-1.ano-1 kg.ha-1.ano-1

Floresta Amazônica Total N P

Biomassa de raízes

Conceitos e constituintes da MOS

Conceitos e constituintes da MOS

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Comparação entre biomassa aérea, raízes e produção de grãos

Mg ha-1Cultura

Soja

CaupiMilhoArroz sequeiro

Raízes finas

1,39

1,030,970,62

Biomassa aérea

2,76

1,703,031,60

Grãos

1,80

0,863,001,41

Conceitos e constituintes da MOS

Conceitos e constituintes da MOS

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 Metodologias de amostragem

Placa com Pregoscomprimento das raízesdiâmetro das raízes

Filmagem no Perfil do Solocomprimento das raízesdistribuição do sistema radicular no perfil do solo

 Monolito

densidade de massa (MS raízes/ Vol. terra coletada)comprimento (compr. raízes / Vol. terra coletada)

Conceitos e constituintes da MOS

Conceitos e constituintes da MOS

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 Método da placa com pregos no campo

Conceitos e constituintes da MOS

Conceitos e constituintes da MOS

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 Método da placa com pregos no campo

Conceitos e constituintes da MOS

Conceitos e constituintes da MOS

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 Prepação do perfil do solo no campo

Conceitos e constituintes da MOS

Conceitos e constituintes da MOS

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 Filmagem do perfil do solo no campo

Conceitos e constituintes da MOS

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Conceitos e constituintes da MOS

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Metodologias de quantificação e exemplos

 Amostragem

 Análise de imagens

Corel Draw SIARCS

Quantificação

Conceitos e constituintes da MOS

Conceitos e constituintes da MOS

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Raízes obtidas por filmagem da placa com pregos

 Análise das imagens no Corel Draw

Conceitos e constituintes da MOS

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 Método Monolito (raízes grossas derivadas do método da extração)

Imagem binarizadaImagem binarizada

com afinamento

Conceitos e constituintes da MOS

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 Imagens processadas no programa SIARCS

Raízes finas“ scanner”

Raízes grossas“scanner”

2 Características dos compartimentos

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L e g e n d a : C o m p r i m e n t o d e r a í ze s (c m 4 , 0 c m -2  solo)

  0-1,5 1 ,5-3,0 3 ,0-4,5 4 ,5-6,0 6 ,0-7,5 7 ,5-9,0 > 9 ,0

  2 0

  1 0

  0

  3 0

  4 0

  5 0

  6 0

  2 0

  1 0

  0

  3 0

  4 0

  5 0

  6 0

5 0 c m

( a ) T e s t e m u n h a ( b ) G r a d a g e m (c) Calcár io na superf íc ie (CS)

( d ) C a l c á r io + g r a d a g e m ( C G )  (e ) CS + fer t i li zantes  ( f ) CG + fer t i li zantes

   P   R   O   F   U   N   D   I   D   A   D   E

 ,  c  m

   P   R   O   F   U   N   D   I   D   A   D   E ,  c  m

 Distribuição do sistema radicular obtido por filmagem do perfil 

2. Características dos compartimentos

2 Características dos compartimentos

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0 1 2 3 4 5 6 7 8

0-10

10-20

20-40

40-60

   P   R   O   F   U   N   D   I   D   A   D   E ,  c  m

MASSA TOTAL DE RAÍZES, mg cm-3 solo0 1 2 3 4 5 6 7 8

0-10

10-20

20-40

40-60

   P   R   O   F   U   N   D   I   D   A   D   E ,  c  m

DENSIDADE DE RAÍZES, cm cm-3 solo

0 0.5 1 1.5 2 2.5 3 3.5 4 4.5

0-10

10-20

20-40

40-60   P   R   O   F   U   N   D   I   D   A   D   E ,  c  m 

MASSA S ECA DE RAÍZES (MSR), mg cm-3solo0 0.5 1 1.5 2 2.5 3 3.5 4 4.5

0-10

10-20

20-40

40-60   P   R   O   F   U   N   D   I   D   A   D   E ,  c  m 

MSR EM DECOMPOSIÇÃO, mg cm-3 solo

 Legenda:

Testemunha

Gradagem

Calcário na supefície (CS)

Calcário + gradagem (CG)

CS + Fertilizantes

CG + Fertilizantes

D.M.S. Tukey a 5 %

Erro padrão

 C.V. = 12,44 %   C.V. = 22,38 %

  C.V. = 24,93 %C.V. = 26,24 %

  d c

  b a

 Distribuição do sistema radicular obtido no monolito

2. Características dos compartimentos

Conceitos e constituintes da MOS

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Ponta de uma raiz penetrando no solo.

Exsudatos

 Definição: Compostos orgânicos secretados ou liberados pelasuperfície de raízes jovens.

 Rizosfera: porção do solo na vizinhançaimediata das raízes que influencia aabundância e a composição dapopulação microbiana.

Conceitos e constituintes da MOS

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Pêlo da raiz

Solo Classes dematerial

Células da epidermee do córtex alteradas

e invadidas porbactérias

Coifa da raíz

Células de coifa da raíz

Micoorganismoscom mucilages

microbiana e deplantas

Raiz

53

13

3

3

4

1 e 2

Tipos de rizodepósitos

1. Exsudatos simples vazamdas células das plantas parao solo

2. Secreções, lançadas porprocessos metabólicos

3. Mucilagens, compostosorgânicos mais complexos

originados das células daraíz e da degradaçãobacteriana

4. Mucigel , capa gelatinosacomposta de mistura de

mucilagens e argilas5. Lisados, compostos

liberadas durante a digestãode células por bactérias.

10 mm

Conceitos e constituintes da MOS

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Ribose 5,3Arabinose 3,6

Fucose 23,3Galactose 26,2Acidos urônicos 23,1Glicose 8,4

Monose 5,9Xilose 4,2

Composição bioquímica da mucilagem (Exemplo para raiz de milho)

Composto %

Conceitos e constituintes da MOS

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Método de estudo qualitativo

Conceitos e constituintes da MOS

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 Destino no solo

Mucilagem

Secreções

Plantas

Coloides

minerais

orgânicoshumina

Biomassamicrobiana

M

ucigel

CO2

Humificação (humina microbiana)

Mineralização (NH4+, NO3-, ...)assimilação pelas plantas

• Lubri ficante para movimento da raiz no solo• Melhora contacto raiz-solo

Conceitos e constituintes da MOS

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Solo

Raíz

 Interações raíz/solo

Conceitos e constituintes da MOS

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CONSTITUINTES

PluviolixiviadosLiteira, restos de culturas e resíduos orgânicos

Raízes e exsudatosOrganismos do solo

Substâncias não-húmicasSubstâncias húmicas

Conceitos e constituintes da MOS

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Tamanho relativo

Fungos

Bactérias

Ameba

Cabeça denematóide

bacteriófago

Protozoa

cilicata

Núcleo

Parede

celularda raizPelo da

raíz

Actinomicetos

 Macrofauna: 2-20 mm

 Mesofauna: 100 m-2 mm

 Microfauna: < 100 m

Conceitos e constituintes da MOS

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Tesourinha

Ordem Dermaptera

 Larvas de borboleta

Ordem Lepidoptera

 Larva bezouro

Ordem Coleoptera

 Larva de môsca

Ordem Diptera

 Excrementos  Excrementos

Conceitos e constituintes da MOS

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 Excrementos

 Minhoca

 Minhoca

Trato digestivo

Trato digestivo

Classe Oligochaeta

Ordem Megascolecidae

Conceitos e constituintes da MOS

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 Definição: Organismos menores do que 0,1 mm, predominando fungos, bactérias e actinomicetos em número, massa e capacidade metabólica.

Biomassa microbiana do solo

Actinomicetos

ProtozoáriosBactériasFungos Jenkinson et al., 1986

Conceitos e constituintes da MOS

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CONSTITUINTES

PluviolixiviadosLiteira, restos de culturas e resíduos orgânicos

Raízes e exsudatosOrganismos do solo

Substâncias não-húmicasSubstâncias húmicas

Conceitos e constituintes da MOS

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Substâncias não-húmicas Definição: Compostos derivados de carboidratos, aminoácidos e proteínas,lipídeos, compostos fenólicos e lignina provenientes da decomposição derestos vegetais e animais no solo

Celulose

Lipídeos

Quitina

Aminoácidos

Glicina(aminoácido)

Ácidos nucleicos

Carboidratos

Glucose(aldose)

Conceitos e constituintes da MOS

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Microbiológica Bioquímica

Degradação

Carboidratos

Ceras

Resinas

Produtos da degradação

Substâncias não-húmicas

Degradação

Síntese Húmus

Carboidratos Proteínas Ácidos graxos Ceras OutrosResinasLignina Pigmentos

Conceitos e constituintes da MOS

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CONSTITUINTES

PluviolixiviadosLiteira, restos de culturas e resíduos orgânicos

Raízes e exsudatosOrganismos do solo

Substâncias não-húmicas

Substâncias húmicas

Conceitos e constituintes da MOS

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Definição (antiga) de substâncias húmicasMistura complexa e recalcitrante de substâncias orgânicas amorfas e coloidais decor marrom ou marrom escuro, modificadas a partir de tecidos orgânicos ou demateriais orgânicos pelos organismos do solo (Stevenson, 1994)

Conceitos e constituintes da MOS

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Associação de moléculas de pequeno tamanho estabilizadas por forças fracas: interaçõeshidrofóbicas, pontes de H eforças de van der Waals

(Piccolo, 1996)

Cátions metálicos

 polissacarídeos

 polipeptídios

cadeias alifáticas

fragmentos aromáticos

Simpson et al., 2002

Modelo Supramolecular