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MÉ
BIOFÍSICA DI
3ª EDIÇÃO
CA
J. J. PEDROSO DE LIMA
IMPRENSA DA
UNIVERSIDADE
DE COIMBRA
COIMBRA
UNIVERSITY
PRESS
(Página deixada propositadamente em branco)
E N S I N O
EDIÇÃO
Imprensa da Universidade de Coimbra
Email: imprensa@uc.pt
URL: http//www.uc.pt/imprensa_uc
Vendas online: http://livrariadaimprensa.uc.pt
COORDENAÇÃO EDITORIAL
Imprensa da Universidade de Coimbra
CONCEÇÃO GRÁFICA
António Barros
INFOGRAF IA
J. J. Pedroso de Lima
I NFOGRAF IA DA CAPA
Mickael Silva
E XECUÇÃO GRÁF ICA

Gráfica de Coimbra
I SB N
978-989-26-0653-8
ISBN DIGITAL
978-989-26-0654-5
DOI
http://dx.doi.org/10.14195/978-989-26-0654-5
D EPÓ S ITO LEGAL

373056/14
© M ARÇO 2014, I M PREN S A DA U NIVER S IDADE DE C OI MB RA


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2
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3
- Outra maneira de dizer é que aumenta a entropia do sistema. Ambas as formas
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de potencial eléctrico - campo eléctrico (força/carga); o gradiente de energia potencial de
pressão - forças de pressão; o gradiente de energia potencial gravitacional - a força gravítica;
o gradiente de potencial químico pode ser visto como mais um caso.
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C A P Í T U L O II
x
GASES
CAPÍTULO II x GASES
II.1 - Introdução
Os gases são muito sensíveis a variações de pressão e temperatura, são

compressíveis dentro de uma grande escala de valores e são infinitamente
expansíveis, preenchendo um recipiente, quaisquer que sejam as suas
dimensões e forma, não apresentando superfície limite própria.
Os gases são constituintes de todos os organismos vivos e desem-
penham um papel de primeira importância em fenómenos essenciais à vida,
tais como a respiração e a nutrição. As leis que regem o seu comportamento
são fundamentais em áreas da Medicina como a pneumologia, a anestesio-
logia e as medicinas do desporto e do ambiente.
São trocados com o exterior, por dia, num organismo humano, um maior
número de moles no estado gasoso (N2, O2, CO2, H2O, etc.) do que nos outros
estados, em conjunto. Esta situação cria uma grande dependência do homem do
meio gasoso que o rodeia e obrigou a Medicina a novas perspectivas, perante
modificações deste meio, introduzidas no mundo actual.
Existe forte evidência para se admitir que no estado gasoso, as moléculas
se encontram muito afastadas umas das outras e se deslocam livremente no
espaço, com uma velocidade média que depende da temperatura. Numa primeira
aproximação podemos considerar que, para baixas pressões, as moléculas de um
gás não exercem quaisquer acções entre si, à excepção de colisões1. As colisões
são tanto mais frequentes quanto maior é a pressão a que um gás se encontra.
Esta aproximação só tem interesse em casos especiais pois a existência
de forças de coesão nos gases é evidenciada em múltiplas e importantes
situações.
Um gás oferece resistência quando se comprime, compensando com
forças de pressão as forças externas exercidas. Se comprimirmos um gás
sem permitir que qualquer energia térmica seja trocada, para além do
aumento da sua pressão, também a temperatura aumenta, já que vai _____
aumentar o número de colisões. Por outro lado, se o gás aquece, as suas
moléculas movem-se, em média, com maior velocidade.
Uma situação prática de aplicação do que referimos no parágrafo
anterior e com as aproximações que se impõem, é o aumento de temperatura
1
- Esta aproximação só tem interesse em casos especiais pois a existência de forças
de coesão nos gases, apesar de pequenas quando comparadas com os líquidos, é
evidenciada em múltiplas situações.
que se verifica no ar que se movimenta do alto de montanhas para o vale

que estas formam.
Pareceu-nos de interesse recordar as leis dos gases.
II.2 - Lei de Boyle
Uma massa de gás, a temperatura constante, ocupa um volume que

varia inversamente com a pressão que suporta.
Podemos então escrever
1
V=KP
ou
PV=K II.1
onde V é o volume ocupado pelo gás, P a pressão a que está sujeito e K uma
constante.
Esta lei é facilmente verificada com o dispositivo da Fig. II.1, em que
os dois recipientes de vidro A e B comunicam por um tubo maleável.
O recipiente A é fechado e está graduado, de modo a conhecer-se o
volume V de gás aí existente acima da superfície do mercúrio, representado
a cheio na figura. O recipiente B é aberto.
_____
17
Fig. II.1 - Dispositivo para verificação da lei de Boyle.

Suponhamos que o desnível entre as superfícies livres do mercúrio em

A e B é h1 e que nestas condições o volume ocupado pelo gás é V1.
A pressão exercida sobre o gás é
P1=Po+h1Ug II.2
onde Po é a pressão atmosférica, U a massa específica do mercúrio e g a ace-

leração da gravidade.
Para outros desníveis h2, h3, ... entre as superfícies livres em A e B
corresponderiam volumes de gás V2, V3, ... e pressões P2, P3, ... ,
respectivamente.
Se a temperatura foi mantida constante durante a experiência, a
relação
P1 V1 = P2 V2 = P3 V3 = ... II.3
verifica-se com bom grau de aproximação.

Como, por definição, a massa específica de uma dada massa de gás
varia inversamente com o volume ocupado pelo gás, podemos enunciar a lei
de Boyle numa outra forma: a temperatura constante, a massa específica de
um gás é directamente proporcional à pressão exercida sobre o gás.
A maneira mais usual de exprimir graficamente a lei de Boyle é
colocando produtos PV em ordenada e P em abcissas. Quando a lei se veri-
fica, obtém-se uma recta paralela às abcissas, Fig. II.2.b). Dado que a tem-
peratura se mantém constante, a curva é uma isotérmica. Num gráfico P(V)
as isotérmicas de Boyle são hipérboles equiláteras referidas aos eixos, Fig.
II.2.a).
____
17
Fig. II.2 - Isotérmicas para gás ideal (lei de Boyle). a) Representação P(V) e
b) Representação PV(P).
II.3 - Lei de Charles
Se a pressão se mantiver constante, uma certa massa de gás aumenta

de 1/273 do seu volume a 0 ºC por cada grau centígrado que a temperatura
aumenta, ou seja,
t
V = Vo (1 + 273 ) II.4
onde V é o volume a t graus centígrados e Vo o volume a 0 ºC.
Fazendo um gráfico de V em função de t obtém-se uma linha recta

que intercepta o eixo das temperaturas (ponto para o qual V = 0) para t = –
273 ºC. No caso ideal de se terem gases à temperatura de –273 ºC, se a lei de
Charles ainda fosse aplicável, o volume por eles ocupado deveria ser nulo. A
temperatura de –273 ºC corresponde ao zero absoluto, ou seja, o zero da
escala de temperatura absolutas (escala Kelvin) e é a mais baixa temperatura
que, teoricamente, poderá ser atingida. A variação de temperatura corres-
pondente a um grau Kelvin é igual à correspondente a um grau centígrado e
a conversão de temperaturas entre as duas escalas obtém-se pela relação
simples
T = t + 273 II.5
onde T significa temperatura em graus Kelvin e t temperatura em graus cen-

tígrados.
A equação II.4 pode então escrever-se na forma
_____
V = K' T II.6
17
onde K' é uma constante.
A lei de Charles pode pois enunciar-se de modo diferente:

O volume de uma dada massa de gás, a pressão constante, é directa-
mente proporcional à temperatura absoluta a que o gás se encontra.
A lei de Charles podia verificar-se utilizando o dispositivo represen-

tado na Fig. II.3
Fig. II.3 - Dispositivo para verificação da lei de Charles.
Uma massa de gás m ocupa o volume Vo no recipiente A (Fig. II.3-a),

à temperatura de 0 ºC sob a pressão Po, que na figura é a pressão atmos-
férica, visto o mercúrio ter o mesmo nível nos dois recipientes A e B. Aque-
çamos o gás, elevando a sua temperatura de t ºC. Para manter constante a
pressão, teremos de baixar B até se ter o mercúrio ao mesmo nível nos dois
ramos (Fig. II.3-b). O gás, agora à temperatura t, é sujeito à mesma pressão,
ocupa o volume Vt que deverá obedecer à relação II.4. Aplicando esta
expressão para duas temperaturas t e t' obtemos:
t
Vt = Vo (1+ 273 ) II.7
_____
17
t´
V´t = Vo (1+ 273)´ II.8
Subtraindo membro a membro obtém-se:

Vo
'V = 273 't II.9
Podemos dizer pois que, a pressão constante, a variação relativa de

volume é uma função linear da variação de temperatura. Esta é uma outra
forma de enunciar a lei de Charles.
Podemos ainda ver qual será a pressão a que o gás, à temperatura t,
deve ser sujeito para que o volume por ele ocupado seja Vo, o volume antes
do aquecimento.
Elevando B até se ter em A o mercúrio ao nível correspondente ao
volume Vo (Fig. II.3-c) a pressão a que se sujeita o gás será:
Pt = Po + h g U ,,
Como a temperatura se manteve constante nas duas últimas experiên-

cias, podemos usar a lei de Boyle e escrever
t
Pt Vo = Po Vt = Po Vo (1 + 273) II.11
Dividindo por Vo e igualando primeiro e último membros da Eq. II.11 vem
t
Pt = Po (1 + 273) II.12
Po
Pt = 273
= K" T II.13
Usando a lei de Boyle, chegámos a uma outra forma da lei de Charles:
a volume constante, a pressão a que uma dada massa de gás é sujeita, varia
directamente com a temperatura absoluta.
_____ Quando aplicadas a gases reais, as leis de Boyle e Charles só se
17 verificam aproximadamente e, em geral, para valores de pressão pouco ele-
vados e temperaturas não muito baixas. Dentro dos limites de pressão e tem-
peratura em aquelas leis se verificam o comportamento de um gás diz-se
ideal.
Suponhamos que, usando as condições da Fig. II.3 e mantendo a
temperatura t constante se varia a posição da coluna B, de modo a obter-se
um volume qualquer V, a que corresponde uma pressão P.
Pela lei de Boyle será

t
PV = Pt Vo = Po Vt = Po Vo (1+ 273) II.14
ou seja,
PoVo
PV = 273 T
PV = K1T II.15
Esta expressão é uma primeira forma da equação dos gases perfeitos.

A constante K1 depende da massa e da composição química do gás.
Condições normais - visto o volume de um gás ser tão marcadamente

afectado por variações de temperatura e pressão é necessário, ao comparar
diferentes volumes de gases, que eles estejam nas mesmas condições de
pressão e temperatura. As condições normais de pressão e temperatura, às
quais os volumes de gases são reduzidos para fins de comparação são:
Pressão - 760 mm Hg (1 atmosfera)

Temperatura – 0 ºC (273 ºK).
Estas condições costumam indicar-se abreviadamente por condições

P.T.N..
II.4 - Lei de Avogadro
Volumes iguais de gases nas mesmas condições de pressão e tempera-

tura contêm o mesmo número de moléculas. _____
17
Esta lei, assim como as anteriormente referidas, é verdadeira unica-
mente para gases ideais.
A lei de Avogadro permite obter a equação dos gases perfeitos numa
forma geral.
A molécula-grama de diferentes substâncias contém o mesmo número
de moléculas (Número de Avogadro = 6,0225u1023 moléculas).
Da lei de Avogadro conclui-se pois que, nas mesmas condições de
pressão e temperatura, moléculas-grama de diferentes gases ocupam o
mesmo volume. Este volume é 22414 cm3 nas condições P.T.N. para 1 mole
de qualquer gás (1 mole de O2 = 32 g, 1 mole de H2 = 2,016 g, etc.).
Aplicando a Eq. II.15 para 1 mole de um gás nas condições P.T.N., já
que P = 760 mm Hg = 1,0133u106 dyn/cm2, V = 22414 cm3 e T = 273 ºK,
temos que K1 = 8,314u107 erg/mole ºK.
É evidente que K1 tem o mesmo valor para 1 mole de qualquer gás
nas condições P.T.N. e designa-se por R ou constante dos gases.
Para 1 mole de qualquer gás tem-se pois
PV = RT
Para um número n de moles, à pressão P, temperatura T e ocupando

um volume V a equação II.9 pode escrever-se
PV = nRT II.16
que é a equação dos gases perfeitos na forma geral.
II.4.1 - Significado da constante R
Suponhamos que se eleva a temperatura de 1 mole de gás de T oK a

(T+1) oK fazendo variar o volume de V a V´ à pressão constante P.
Aplicando a equação dos gases perfeitos antes e depois da variação de
temperatura, temos:
_____
17 PV = RT e PV' = R(T + 1) II.17
Subtraindo a primeira expressão da segunda tem-se
P (V' - V) = R II.18
O primeiro membro da Eq. II.18 representa energia ou trabalho.
Consideremos um cilindro munido de um êmbolo e contendo um

volume V de gás à pressão P contrabalançada pela Força F exercida sobre a

superfície S do êmbolo. Se o embolo se deslocar contra a força F, de um
comprimento x, devido a um aumento de temperatura do gás, o trabalho
realizado pela força F é negativo e dado por
W' = - Fx = - PSx II.19
= - P (V' - V) II.20
onde V' é o volume do gás após o aumento da temperatura. Como F se man-

teve constante, a pressão do gás após o aumento de temperatura ainda é P.
O trabalho realizado pelo gás é igual e de sinal contrário ao realizado
pela força F, ou seja,
W = P (V'-V) II. 21
A expressão de R é idêntica à Eq. II.21. Dadas as condições usadas

para deduzir a Eq. II.18 pode dizer-se que R é igual ao trabalho produzido
por um mole de gás ideal, expandindo-se contra uma pressão constante
como resultado de um aumento de temperatura de 1 ºK.
Fica evidente que, na expressão geral PV = nRT o produto PV representa
a quantidade de trabalho produzida por n moles de um gás ideal na expansão
contra uma pressão constante, quando a temperatura aumenta de 0 a t ºK.
II.5 - A teoria cinético-molecular dos gases

_____
Esta teoria surgiu como uma tentativa de explicação dos fenómenos 17
observados que levaram à descoberta das leis dos gases atrás descritas. Os
postulados mais importantes desta teoria são:
1 - Os gases são constituídos por partículas discretas, as moléculas; as
moléculas de um dado gás são idênticas entre si.
2 - Num gás as moléculas encontram-se muito afastadas e o volume
molecular total é desprezível em comparação com o volume do
recipiente (se a pressão não for muito elevada).
3 - As moléculas estão em movimento estatístico. Deslocam-se em
linha recta com velocidade que é proporcional à raiz quadrada da
temperatura absoluta do gás.

4 - As moléculas são perfeitamente elásticas, mudando de direcção
sem perda de velocidade após colisões entre si ou com as paredes
dos recipientes.
5 - As moléculas não exercem força atractiva entre si (trata-se de uma
aproximação nem sempre válida).
6 - A pressão do gás é o resultado de impactos das moléculas nas pa-
redes dos recipientes.
Aplicando as leis da mecânica a gases monoatómicos e supondo que a

energia interna do gás é unicamente energia cinética, devida ao movimento
estatístico das moléculas (movimento de translação), é possível chegar-se às
expressões das leis dos gases reais.
A teoria cinética permitiu, por exemplo, provar que a energia cinética
total de 1 mole de gás é
Energia cinética/mole = 3/2 PV II.22
Esta expressão conduz a um conceito molecular de temperatura.

Vimos que experimentalmente se prova a relação
PV = nRT
Então para 1 mole podemos escrever, usando as equações II.16 e II.22
Energia cinética/mole = 3/2RT II.23
A constante dos gases R é uma constante de proporcionalidade entre

_____ energia cinética e temperatura absoluta. A temperatura é então uma medida
1
da energia cinética média das moléculas.
Não esqueçamos que estes conceitos se aplicam a gases ideais onde
admitimos não existir energia potencial de interacção entre as moléculas.
II.6 - Desvios às leis dos gases
Os gases reais só aproximadamente seguem as leis dos gases. Consi-

deremos o caso da lei de Boyle. Vimos que a representação gráfica dos

produtos PV em função de P, a temperatura constante, num gás ideal fornece
uma linha recta paralela às abcissas.
A Fig. II.4 mostra algumas curvas obtidas na prática.
Fig. II.4 - Isotérmicas para gases reais. Desvios à lei de Boyle. Unidades no S.I.
A maioria dos gases tem um comportamento próximo ao previsto pela

lei de Boyle para baixas pressões (abaixo de 10 atmosferas) mas quanto
mais próximo o gás se encontra das condições que o levam à liquefacção,
maior é o desvio, como se observa para o CO2.
De acordo com a lei de Boyle, um gráfico da pressão em função do
volume para uma certa massa de um gás deve dar uma hipérbole equilátera
referida aos eixos, havendo uma destas curvas para cada valor de tempera-
tura. As curvas P(V) com T como parâmetro, são chamadas isotérmicas e na
Fig. II.5 mostram-se isotérmicas de um gás real.
_____
1
Fig II.5 - Isotérmicas de um gás real.

A curva para T = T4 corresponde a um comportamento próximo do gás

perfeito, de acordo com a lei de Boyle. Na segunda isotérmica, para t = T3,
apesar de se verifica um certo afastamento da hipérbole equilátera ideal, o gás
não sofre qualquer mudança de estado para os correspondentes valores de P e V.
A curva para a temperatura T2 representa uma situação diferente. Para
pressões baixas e à temperatura constante da isotérmica, um aumento
gradual de pressão conduz a uma diminuição de volume até o ponto a ser
atingido. Neste ponto o volume decresce praticamente sem aumento da pres-
são, liquefazendo-se parte do gás até no ponto b a liquefacção é completa.
A isotérmica a partir de b sobe quase verticalmente pois os líquidos
são praticamente incompressíveis. A isotérmica para T = T ct é chamada a i-
sotérmica crítica pois marca a separação entre as isotérmicas onde não
ocorre liquefacção, tais como T3 e T4 e isotérmicas onde ocorre liquefacção,
tais como T1 e T2. Nas áreas riscadas coexistem as fases líquida e gasosa.
Temperatura crítica Tct é aquela acima da qual não é possível a li-
quefacção do gás qualquer que seja a pressão.
Pressão crítica Pct é a mínima pressão necessária para a liquefacção
do gás à temperatura crítica. Volume crítico é o volume ocupado pelo massa
de gás em estudo à temperatura e pressão críticas. A curva a tracejado limita
a região onde existe equilíbrio entre o gás e a sua fase líquida.
Um gás a uma temperatura abaixo da temperatura crítica designa-se
por vapor. A pressão correspondente à linha ab ou a'b' é a pressão do vapor
saturado à temperatura correspondente. A pressão do vapor saturado é a
pressão máxima que pode ser exercida pelo vapor a essa temperatura e
depende exclusivamente da temperatura. O vapor de água no ar atmosférico
a uma dada temperatura tem geralmente uma pressão parcial menor do que a
pressão do vapor saturado para a mesma temperatura. A relação entre a
pressão do ar e a pressão do vapor saturado para a mesma temperatura é, co-
_____ mo sabemos, a humidade relativa do ar.
18
A transformação inversa da liquefacção é a vaporização. Quando a
vaporização se efectua numa atmosfera gasosa chama-se evaporação. As
moléculas do líquido abandonam a superfície líquida até a pressão parcial
do vapor na atmosfera que contacta o líquido ser igual à tensão do vapor sa-
turado ou tensão máxima do vapor à temperatura considerada. Portanto, se a
atmosfera for ilimitada, a evaporação poderá ser total.
Chama-se calor latente de vaporização a uma dada temperatura, a
quantidade de calor que é necessário fornecer a 1 g de líquido para o vapo-
rizar a essa temperatura. Esta quantidade de calor é igual, em valor absoluto,

ao calor latente de condensação ou liquefacção.
Da observação da figura anterior conclui-se que os desvios do com-
portamento ideal apresentados pelos gases reais são tanto maiores quanto
mais próximos das condições críticas o gás se encontra.
Os desvios às leis dos gases, apresentados pelos gases reais, podem
explicar-se com base na teoria cinética. Nesta admite-se que não existe ener-
gia potencial de interacção entre as moléculas. Existem, no entanto, forças
conhecidas como forças de Van der Waals, que actuam quando as moléculas
se encontram próximas.
Para pequenas distâncias entre as moléculas e a temperaturas tais que
o movimento térmico não contrabalance as forças de Van der Waals, estas
opõem-se ao movimento das moléculas que vão chocar contra as paredes
dos recipientes ficando a pressão do gás menor do que a prevista pela lei de
Boyle. Então
PV < RT II.24
Uma outra suposição, é a do volume total ocupado pelas moléculas,

ser desprezível em função do volume do recipiente. Esta hipótese é susten-
tável quando as moléculas se encontram muito separadas mas à medida que
a pressão do gás aumenta, aquele volume vai-se tornando mais significativo.
Nestas condições, o efeito do volume próprio das moléculas faz diminuir
consideravelmente o espaço disponível para o movimento térmico.
O volume que se observa na prática é, portanto, superior ao
considerado nas condições ideais da lei de Boyle e se esse efeito for superior
ao devido às forças de Van der Waals (Eq. II.24), vem
PV > RT II.25 _____

18
A equação de Boyle não é válida se coexistem duas ou mais fases.
Na Fig. II.5 estão representadas as curvas P(V) para gases reais a
diferentes temperaturas. Para a temperatura T4 a curva é uma hipérbole, tra-
duzindo a equação de Boyle-Mariotte. Para a temperatura Tct (temperatura
crítica) ocorre um intervalo de volumes para o qual não há variação de
pressão. Isto só é possível se ocorrer liquefacção do gás, coexistindo as fases
gás e líquido. A temperatura crítica é aquela acima da qual não pode ocorrer
liquefacção, qualquer que seja a pressão. Pressão crítica Pct é o mínimo
valor para o qual ocorre liquefacção à temperatura crítica.

Mesmo com pressões muito elevadas não é possível liquefazer a
maioria dos gases à temperatura ambiente. A excepção é o CO2.
Para liquefazer um gás é necessário arrefecê-lo até temperaturas
abaixo de Tct e elevar a pressão acima de Pct, a pressão crítica. Alguns
valores destes parâmetros são referidos no quadro II.1.
4XDGUR,,9DORUHVGH3FWH7FWSDUDDOJXQVJDVHV

gás Tct (°C) Pct (atm)

He -267,9 2,26
H2 -239,9 12,8
N2 -147,1 34
O2 -118,8 48
CO2 31,1 73
NH3 132,4 111,5
H2O(vapor) 374,0 217,7
II.7 - A equação de Van der Waals
Esta equação foi sugerida para compensar os efeitos verificados na

relação pressão-volume nos gases, de modo a tornar a equação de Boyle
aplicável a gases reais. Esta equação tem a forma para n moles
_____ a n2
(P+ V2 ) (V-nb) = nRT II.26
18
RQGH 3 p D SUHVVmRGR JiV 9 p R YROXPH GH JiV j SUHVVmR 3 H WHPSHUDWXUD

FRQVWDQWH H D H E VmR FRQVWDQWHV SDUD XP GDGR JiV H YDULDP SDUD RVGLIHUHQ
WHVJDVHV(VWDVFRQVWDQWHVWrPGHVHUGHWHUPLQDGDVH[SHULPHQWDOPHQWH
a n2
O termo V2 destina-se a compensar as forças de Van der Waals que
tendem a diminuir a pressão observada em relação à pressão ideal. A relação
n/V é o número de moles por unidade de volume.
É também tomado em consideração o volume total ocupado pelas

moléculas, de valor finito, que diminui o volume do espaço disponível para
os movimentos das moléculas do gás. É subtraído ao volume do JiV um
valor b, por mole, TXH p XPD IXQomR GR GLkPHWUR HVIpULFR GD PROpFXOD GR
JiVFRQKHFLGRFRPRGLkPHWURGHYDQGHU:DDOV
1R TXDGUR Q,, HVWmR DOJXQV YDORUHV SDUD D H E GH DOJXQV JDVHV 2
KLGURJpQLR H KpOLR DSUHVHQWDP YDORUHV SUy[LPRV GH ]HUR 2 VHX
FRPSRUWDPHQWR p SUy[LPR GR LGHDO RX VHMD VmR PtQLPDV DV LQWHUDFo}HV
PROHFXODUHVTXHDSUHVHQWDPQRHVWDGRJDVRVR

4XDGURQ,,9DORUHVGHDHE equação de Van der Waals)SDUDDOJXQVJDVHV

*iV D E
+
2
1
&2
&O
+H
Considerando como hipótese que as forças entre as partículas são nulas e

que o volume intrínseco é zero, a lei de Van der Waals para os gases reais _____
converte-se na equação de Boyle-Mariotte dos gases ideais (PV = Const.). 18
II.8 - Difusão dos gases
Como sabemos a difusão é o processo pelo qual tendem a desaparecer

os gradientes de concentração de substâncias (cujas moléculas estão sujeitas
a movimento térmico estatístico de translação) contidas num sistema, no
qual não existam obstáculos completos à sua passagem.
A difusão ocorre como resultado do transporte de moléculas de
regiões de maior concentração para regiões de menor concentração e pode

ocorrer em qualquer dos estados de agregação material.
Quando recipientes com dois gases diferentes são postos em comuni-
cação sem agitação externa, misturam-se progressivamente até a concen-
tração de qualquer dos gases ser constante em todos os pontos do sistema
formado pelos dois recipientes.
A velocidade de difusão depende da natureza dos gases, da pressão e
da temperatura. Quando a difusão de um gás se efectua através de uma
membrana porosa, a velocidade de difusão é proporcional à diferença de
pressão entre os dois lados da membrana e inversamente proporcional à raiz
quadrada da densidade do gás.
A proporcionalidade inversa entre a velocidade de difusão e a raiz
quadrada da densidade constitui a lei de Graham. Este provou serem os tem-
pos de difusão de volumes iguais de gases nas mesmas condições de pressão
e temperatura, através de pequenos orifícios, directamente proporcionais às
raízes quadradas das densidades.
Como as densidades dos gases são proporcionais ao seu peso molecu-
lar, segue-se que a velocidade de difusão de um gás é também inversamente
proporcional à raiz quadrada do seu peso molecular.
Evaporação e condensação podem considerar-se casos particulares da
difusão de um gás através de uma membrana (a superfície livre do líquido).
A massa de líquido que passa do estado líquido ao gasoso, a uma dada tem-
peratura, por unidade de tempo, devida à evaporação, é dado por
dm p -p
=KS s II.27
dt p
onde S é a área da superfície livre do líquido, p a pressão parcial do vapor

_____ do líquido na mistura gasosa em contacto com o líquido, ps a pressão do va-
18 por saturado do líquido à temperatura da experiência e K uma constante.
Ocorrerá evaporação sempre que (ps - p) seja maior que zero. Se (ps - p) se
anula a mistura gasosa em contacto com o líquido está saturada do vapor do
líquido. A condensação consiste na passagem de moléculas da fase gasosa
para a fase líquida em situações em que (ps - p) < 0.
Uma situação biológica onde a difusão gasosa desempenha um impor-
tante papel é o processo mecânico da respiração pulmonar. Não é a corrente
de arrastamento devido à diferença de pressão entre o ar exterior e o ar al-
veolar que arrasta o oxigénio necessário para o espaço alveolar e que
remove deste o CO2 mas difusão molecular devido às diferenças de concen-

tração destes gases ente os alvéolos e a mistura gasosa nas proximidades dos
alvéolos. Em cada ponto das vias aéreas terminais as variações das con-
wCi
centrações dos gases, Ci com o tempo são função do coeficiente de
wt
difusão D, da área da secção recta S da via aérea e da distância percorrida
pelas moléculas x.
wCi wC2 1 wC wS
= D ( 2i + S i ) II.28
wt wx wx wx
que é uma equação mais geral do que a 2ª lei de Fick pois entra em conta
com as variações de S com a distância.
II.9 - Lei de Dalton das pressões parciais
A Lei de Dalton das pressões parciais diz que quando dois ou mais
gases que não reagem quimicamente entre si são misturados, cada gás se
comporta independentemente dos outros e tem a mesma pressão que teria se
estivesse ele unicamente no volume ocupado pela mistura gasosa. Esta
pressão é a pressão parcial do gás na mistura (por vezes a designação de
tensão é utilizada como sinónimo de pressão parcial).
Por exemplo, quando uma mistura gasosa se encontra em contacto
com um líquido, cada gás dissolve-se como se estivesse só e com a res-
pectiva pressão parcial.
A lei de Dalton é também uma lei ideal que é seguida aproximada-
mente pelos gases reais. _____
Numa mistura com n gases que não reajam quimicamente entre si, se 18
as pressões parciais dos gases da mistura forem
P1, P2, P3, .............., Pn
a pressão total na mistura é igual à soma das pressões parciais dos gases
componentes
P = P1 + P2 + P3 + ............. + Pn II.29
A pressão parcial de qualquer gás em mistura pode ser calculada se

conhecermos o número de moles de cada um dos gases da mistura. Su-
ponhamos que a mistura, ocupando um volume V, contém n1 moles do gás
1, n2 moles do gás 2 ....e nn moles do gás n.
Supondo a temperatura T constante, pela lei de Dalton, podemos
aplicar a lei dos gases perfeitos a cada gás da mistura, ou seja
P1V = n1RT
P2V = n2RT
.
.
.
PnV = nnRT II.30
Somando as equações anteriores membro a membro, vem
(P1 + P2 + P3 .....+ Pn) V = (n1 + n2 + n3 + .....+ nn) RT
Chamando
N = n1 + n2 + n3 + ...... + nn
e como
P = P1 + P2 + P3 + ..... + Pn
vem
PV = NRT II.31
Dividindo cada uma das Eqs. II.18 pela Eq. II.19 vem
n1
_____ P1 = n + n +.....+ n P
1 2 n
18
n2
P2 = n + n +.....+ n P II.32
1 2 n
.
.
.
nn
Pn = n + n +.....+ n P
1 2 n
ni n
A expressão n + n +.....+ n = Ni = fi é a fracção molar do gás i
1 2 n
na mistura.
Então as pressões parciais dos gases da mistura são
n1
P1 = N P = f1
n2
P2 = N P = f2 P
.
.
nn
e Pn= N P = fnP II.33
A pressão parcial de cada um dos gases da mistura é o produto da

fracção molar respectiva pela pressão total da mistura.
Como para qualquer gás i da mistura é
ni pi
N = P = fi II.34
podemos dizer que a fracção molar de cada um dos elementos de mistura, ou

seja, a fracção do número total de moles que corresponde a esse gás é
também a fracção da pressão total que lhe corresponde.
É muitas vezes conveniente relacionar a pressão parcial de um gás
_____
com a sua concentração molar. Dividindo as Eq. II.18 por V obtém-se,
18
exprimindo genericamente
Pi= CiRT II.35

ni
onde a concentração molar Ci = V , é expressa em moles por unidade de vo-
lume. A Eq. II.35 mostra que, a uma dada temperatura, a pressão parcial de
um gás numa mistura é proporcional à sua concentração molar na mistura.
É também possível exprimir a fracção molar como uma relação entre
volumes. Consideremos que os n1 moles do gás 1, da situação anterior, eram
retirados da mistura e sujeitos a uma pressão P igual à pressão total da mis-

tura inicial e à mesma temperatura T, ocupando agora o volume V1. Então,
pela lei dos gases perfeitos podemos escrever
PV1 = n1RT
Fazendo de igual modo para os outros gases, os n2 moles do gás 2 à

temperatura T e à pressão P ocupam o volume V2, ..., os nn moles do gás 2 à
temperatura T e à pressão P ocupam o volume Vn, vindo então
PV2=n2RT II.36
.
.
.
PVn = nnRT
Somando as equações anteriores membro a membro, vem
P (V1 + V2 + V3 + ... + Vn) = (n1 + n2 + n3 + ... + nn) RT
= NRT
Por analogia com o caso anterior terá de ser
V1 + V2 + V3 + ... + Vn = V
Ou seja,
_____ PV = NRT II.37

1
Dividindo cada uma das Eqs. II.23 pela Eq. II.24 vem
V1 n1
V = N = f1
V2 n
= 2 = f2 II.38
V N
.
e
Vn n
= n = fn
V N
A fracção molar de cada um dos gases é também a relação entre o

volume ocupado por cada um dos gases à pressão total e o volume da
mistura à mesma pressão.
II.10 - Solubilidade dos gases nos líquidos
Os gases dissolvem-se em menor ou maior quantidade nos líquidos

com os quais são postos em contacto. Após um certo tempo, a concentração
do gás dissolvido num líquido, em presença de uma atmosfera contendo o
gás, atinge um valor de equilíbrio, que não é ultrapassado se a temperatura e
a pressão do gás não forem alteradas. Diz-se então que para as condições
existentes o líquido se encontra saturado de gás.
A solubilidade dos gases em água ou outros líquidos varia marcada-
mente com a natureza dos gases em contacto.
Quando um gás não reage quimicamente com o líquido, a sua solubi-
lidade é em geral muito pequena. Por exemplo, o oxigénio dissolve-se muito
pouco em água, enquanto que o cloro ou amoníaco que reagem com a água,
apresentam alta solubilidade naquele líquido.
A solubilidade dos gases em líquidos obedece à lei que a seguir men-
cionaremos.
_____
II.11 - Lei de Henry
1
Diz a Lei de Henry: a uma temperatura determinada, a concentração

C de um gás dissolvido num líquido, após ter sido atingido o equilíbrio, é
proporcional à pressão parcial P que o gás exerce à superfície do líquido.
C = KsP II.39
Ks é o coeficiente de solubilidade do gás à temperatura considerada.

A Eq. II.39 verifica-se nas chamadas soluções simples, para as quais

Ks é constante.
A concentração C pode ser expressa pelo volume de gás medido nas
condições experimentais iniciais/m/ de solvente, em fracção molar do gás
GLVVROYLGR HP PLOLPROHV/ RX P/ GR JiV 371 P/ GR VROYHQWH $
pressão P em mm Hg, Pascais ou atmosferas.
Quando se tem uma mistura de gases a temperatura constante e em
contacto com um líquido e em equilíbrio, a concentração para cada compo-
nente é proporcional à pressão parcial correspondente.
O coeficiente de solubilidade depende das duas naturezas do gás e
líquido e da temperatura.
É importante aqui recordar-se que, quando se tem um gás em contacto
com um liquido, há sempre que contar com a pressão de vapor do líquido.
Quando um gás se encontra em contacto com a água, a fase gasosa não só
contém o gás em questão mas também vapor de água. Portanto, para se de-
terminar a pressão parcial do gás é necessário subtrair da pressão total
externa a pressão do vapor de água à temperatura da experiência.
A maioria dos gases obedecem aproximadamente à lei de Henry para
as baixas pressões correspondentes aos sistemas biológicos, mas muitos dos
gases não conservam a identidade química quando em solução, não sendo
aplicável a Eq. II-39. Por exemplo, no caso do sangue, o oxigénio e o
dióxido de carbono reagem com a hemoglobina e outras proteínas. A lei de
Henry aplica-se aqui unicamente às moléculas que permanecem na forma de
gás dissolvido, sem reagir quimicamente, como é o caso do oxigénio
dissolvido fisicamente no sangue. Para a concentração de um gás que é
retido quimicamente pode considerar-se uma outra relação, esta não linear
_____ C = f(P) II.40

19
A concentração do gás é uma função (crescente) da pressão parcial e
depende de factores condicionantes relativos ao problema em causa.
O coeficiente de solubilidade dos gases nos líquidos diminui com o
aumento da temperatura. Este facto explica-se com o aumento da energia
térmica de agitação, originando uma maior probabilidade de moléculas do
gás abandonarem o líquido sem ocorrer qualquer variação de energia poten-
cial, nas interacções gás-líquido. Observar na tabela seguinte dos
coeficientes de solubilidade do hidrogénio na água.
Temp. ºC 0 4 10 15 2
Coef. Sol (m//m/) 0,0215 0,0206 0,0196 0,0188 0,0182
É, portanto, a baixa temperatura e a alta pressão, que a quantidade de

gás dissolvido num líquido é mais elevada. Contrariamente, quando se
deseja libertar de um líquido o gás nele dissolvido, ou se aquece até à ebuli-
ção ou se põe em contacto com um vazio tão elevado quanto possível.
Em medicina e biologia o coeficiente de solubilidade Ks, atrás defini-
do e usado para exprimir a solubilidade de um gás, aparece mais frequente-
mente em duas formas:
Coeficiente de absorção de Bunzen Dt - é a quantidade de gás em m/,

reduzido às condições P.T.N. (760 mm Hg, 0ºC) que, após completo equilíbrio,
se dissolve em l ml do solvente quando a pressão do gás é 1 atmosfera.
Coeficiente de solubilidade de Ostwald Ot - define-se de maneira

idêntica ao anterior, excepto que o volume de gás dissolvido em m/ não é
reduzido à temperatura normal; só a pressão é reduzida.
Facilmente se relacionam os coeficientes D e O. Sabemos já que o
volume de uma certa massa de gás é proporcional à temperatura absoluta
quando a pressão é mantida constante. Então:
Ot = Dt II.41
onde Ot e Dt representam os coeficientes de Ostwald e Bunsen para a

temperatura t.
A designação de coeficiente de solubilidade de Ostwald é por vezes _____
utilizada para exprimir exclusivamente o coeficiente de solubilidade entre o 19
ar e os tecidos, de acordo com a definição acima dada para O.
A Eq. II.39, permite mostrar que a concentração de gás dissolvido,
medida em m/, à pressão parcial do respectivo componente na mistura exte-
rior, por m/ do solvente é independente da pressão (a temperatura cons-
tante), ou seja, sabendo esta concentração para uma dada pressão parcial
sabemos para todas as outras.
De facto, a massa de gás dissolvido num volume Vs de líquido em
contacto pode sempre escrever-se
M=Vd II.42
onde V é o volume do gás e d a densidade, medidos à mesma pressão P e

temperatura T. Num gás, a densidade é proporcional à pressão, donde
M = V P Cte
Consideremos que P é a pressão parcial externa do gás que tem a

massa M dissolvida no líquido. Então a lei de Henry (Eq. II.39) pode
escrever-se
M V P Cte
= = Ks II.43
Vs Vs
donde
V
= Cte II.44
Vs
A concentração do gás dissolvido, em ml, à pressão parcial do respe-

ctivo componente, por ml do solvente (V/Vs) é uma relação constante para
uma dada temperatura. Este resultado não contraria a lei de Henry, pois
resulta de se usarem unidades de concentração do gás dissolvido (condições
de medida do volume de gás) diferentes para cada valor de pressão.
II.12 - Elementos de física do aparelho respiratório

_____
19 Em termos gerais, a respiração consiste na utilização de oxigénio e na
produção de dióxido de carbono pelas células vivas e nos mecanismos de
troca destes gases, entre as células e a atmosfera.
A grande maioria dos seres vivos recolhe o O2 directamente da atmosfera,
utilizando um dos diversos mecanismos pelos quais este gás pode ser fornecido aos
tecidos metabolizantes. Nos seres inferiores, estes processos são: difusão directa,
troca através da pele, uso de guelras e uso de tubos de ar.
O sentido de transferência das trocas gasosas, nestes processos, é in-
teiramente determinado pelo gradiente de pressão entre a atmosfera e as células.
Nos mamíferos a recolha do O2 atmosférico é feita segundo um es-

quema mais elaborado. O O2 é retido pelo sangue por transferência através
dos capilares pulmonares e distribuído aos tecidos pelo sangue circulante.
No homem de 70 Kg, em repouso, são trocados por minuto, entre sangue e
tecidos, cerca de 300 ml de O2 P.T.N. e, no sentido oposto, cerca de 250 ml
de CO2 P.T.N.. No exercício estes valores podem atingir valores mais de
dez vezes superiores. Numa cidade com dez milhões de habitantes são
retirados da atmosfera por minuto mais de 3×106 litros de O2 P.T.N. só para
a respiração dos seus habitantes.
Como sabemos, a concentração de gás fisicamente dissolvido num lí-
quido, a temperatura constante, é directamente proporcional à pressão parcial do
gás na fase gasosa (lei de Henry da solubilidade dos gases). Quando se tem uma
mistura gasosa em contacto com um líquido, no equilíbrio, o número de
moléculas gasosas que abandonam o líquido por unidade de tempo é igual ao
número das que entram no líquido, e qualquer mudança na pressão parcial do
gás produz uma mudança correspondente no equilíbrio.
Quando ocorre equilíbrio entre um gás, com uma certa pressão parcial,
numa mistura gasosa, e um líquido, pode considerar-se que nas fases gasosa e
líquida, a pressão deste gás é igual. Assim, em condições de equilíbrio, as
pressões parciais do gás nas duas fases dizem-se iguais. Para se determinar a
pressão parcial do gás no líquido, é necessário determinar a composição e
pressão do gás na fase gasosa em equilíbrio.
A pressão parcial do O2 alveolar é superior à pressão parcial do O2 no
sangue enviado aos alvéolos. Contudo, a pressão parcial do CO2 neste sangue é
superior à do ar alveolar.
Deste modo, à medida que o sangue percorre os capilares do pulmão, a
concentração sanguínea em O2 aumenta, sucedendo o contrário à concentração
_____
em CO2. Após as trocas gasosas do pulmão, o sangue é enviado para os tecidos.
19
A pressão parcial de CO2 nos tecidos é superior à do sangue, mas a
concentração em O2 no sangue arterial é maior do que nos tecidos. As trocas
processam-se agora com perda de O2 e ganho de CO2, por parte do sangue.
A Fig. II.6 mostra um modelo esquemático do aparelho respiratório do
homem. As bombas B1 e B2 simulam a acção mecânica dos músculos
respiratórios, que levam à movimentação do ar. As bombas B3 e B4 simulam
a acção mecânica do coração esquerdo e direito que produzem a corrente
sanguínea.
Fig.II.6 - Modelo esquemático do aparelho respiratório do homem.
A pressão parcial do vapor de água no ar alveolar é a tensão do vapor

saturado, que só depende da temperatura e é 47 mmHg a 37 ºC. Conside-
rando a pressão do ar alveolar igual à pressão atmosférica (760 mmHg), vem
PO2 + PCO2 + PN2 + PH2O = 760 mm Hg
PO2 + PCO2 + PN2 = 760 - 47 = 713 mm Hg
A composição em volume do ar é:
O2 = 14% CO2 = 5,6% e N2 = 80,40%
Logo, as fracções molares2 são:
14/100 5,6/100 e 80,4/100.

_____ Então, as pressões parciais na mistura gasosa alveolar são:
19
PO2 = 713 u 14/100 = 99, 82 mmHg
PCO2 = 713 u 5,6/100 = 39,9 mmHg
2
Numa mistura gasosa ocupando V e sendo P1+P2 +...+Pn = P, pela equação dos gases
perfeitos vem P1V = n1RT, P2V = n2RT e PV = (n1+n2+...) RT donde P1/P = n1/N, etc.. Os
volumes dos gases medidos à pressão total P são obtidos por PV 1 = n1RT, PV2 = n2RT e
P(V1 +V2+..) = NRT ou PV = NRT e V 1/V = n1/N = P1/P, etc., então a fracção molar pode
escrever-se como uma relação entre número de moles, entre pressões ou entre volumes.
PN2 = 713 u 80,4/100 = 573,2 mmHg
Estes cálculos justificam os valores de PO2 e PCO2 já referidos na Fig.

II.6. A espessura da parede alveolar é de cerca de 0,5 Pm e a área total de
troca 50-120 m2, embora, no repouso só ~35 m2 sejam utilizados.
Já referimos que as bombas B1 e B2 executam o trabalho mecânico da
respiração, fazendo circular o ar no compartimento 1. Note-se que a compo-
sição do gás alveolar é substancialmente diferente da do ar atmosférico. Não
esquecer que há retenção de CO2 no labirinto alveolar e que temos de contar
com a tensão do vapor de água à temperatura do corpo humano, para uma
pressão total de ar alveolar que é praticamente a pressão atmosférica.
A membrana M separa os compartimentos 1 e 2. Neste último, as
bombas B3 e B4 fazem circular o sangue, produzindo um trabalho que simu-
la o do coração. As trocas gasosas entre o sangue e o ar alveolar processam-
se através da membrana M. Na parede inferior de 2 processam-se as trocas
de O2 e CO2 com os tecidos 3.
II.13 - As trocas e o transporte de gases no organismo
É conhecido que os processos responsáveis pelas trocas e transporte

de gases no organismo são: convecção, difusão e combinação química.
A convecção designa de um modo geral a movimentação de uma
massa gasosa por uma acção motriz exterior ao gás.
Os gases são introduzidos no organismo ou eliminados fundamentalmente
pela ventilação pulmonar em que, alternadamente, ar é bombeado do exterior
para os alvéolos pulmonares e uma mistura gasosa é bombeada dos alvéolos para
o exterior. Trata-se pois de um processo de convecção, ao qual se associa um _____
19
fenómeno de difusão em fase gasosa, na massa do gás mobilizado.
A partir do alvéolo, os gases difundem-se através das membranas que
separam os diversos compartimentos líquidos, dissolvendo-se ou reagindo
no seio desses líquidos.
Os gases que intervém nas trocas respiratórias são fundamentalmente
o oxigénio, o anidrido carbónico (gases fisiológicos) e o azoto (biologica-
mente inerte). O oxigénio recolhido do ar é levado aos tecidos pelo sangue
para ser utilizado pelas células. O anidrido carbónico produzido pelos teci-
dos é eliminado após transporte pelo sangue até aos pulmões.
A solubilidade do oxigénio no sangue à temperatura normal do corpo

é insuficiente para as funções fisiológicas. Na ausência de outros fenóme-
nos, o O2 dissolvido fisicamente no sangue pode ser calculado se conhecer-
mos o coeficiente de Bunsen para o sistema oxigénio-sangue a 37 ºC (cujo
valor é 0,0235) e a pressão parcial do oxigénio no ar alveolar (e sangue
arterial) que é cerca de 100 mm Hg. Então,
100
VO2 = 0,0235 u 760 = 0,003 ml O2/ml sangue
ou seja, 3 ml de O2 (P.T.N.) por litro de sangue. Este valor representa cerca

de 1,5 % do concentração total do O2 sanguíneo.
No caso do CO2 o coeficiente de Bunsen é 0,494 e a pressão parcial
no ar alveolar (e sangue arterial) que é cerca de 40 mm Hg. Então,
40
VCO2 = 0,494 u 760 = 0,026 ml CO2/ml sangue
ou seja, 26 ml de CO2 (P.T.N.) por litro de sangue. Este valor representa

cerca de 5 % da concentração total do CO2 sanguíneo.
Contudo o O2 na forma dissolvida é essencial, pois é a partir desta
fase que se processa a combinação com a hemoglobina. Na reacção que
representa a formação da oxihemoglobina a partir da hemoglobina nas
hemácias
Hb + O2 o
m Hb O2 II.45
o termo O2 é oxigénio dissolvido. Sendo K a constante de equilíbrio da

_____
reacção e sendo a concentração do O2 dissolvido, proporcional à pressão
19
parcial do O2, PO2, podemos escrever
[Hb O2]
= K [O2] = K' PO2 II.46
[Hb]
Se 100 representar a quantidade de hemoglobina total e Y a percenta-

gem de hemoglobina na forma HbO2, podemos escrever
Y
100-Y = K' PO2 II.47
100 K' P
donde Y = 1 + K' P O2 II.48
O2
a concentração de HbO2 (oxihemoglobina) depende portanto da pressão par-

cial do O2.
A função que relaciona [HbO2] com PO2 é, em geral, mais complexa
do que a obtida, pois existem na circulação diversas hemoglobinas com
constantes de dissociação diferentes.
Por outro lado, cada molécula de hemoglobina pode receber até um
máximo de quatro moléculas de oxigénio, havendo constantes de equilíbrio
diferentes para cada reacção.
A quantidade de CO2 dissolvido no sangue pode calcular-se a partir
da pressão parcial e do coeficiente a 37 ºC. O CO2 é cerca de 25 vezes mais
solúvel nos meios biológicos que o O2. No sangue venoso a PCO2 é de cerca
de 46 mm Hg e D 37 ºC = 0,48, logo
46
V = 0,48 760 = 0,029 ml CO2 (P.T.N.) /ml sangue a 37 ºC.
No sangue arterial PCO2 = 40 mm Hg, donde
40
V' = 0,48 = 0,025 ml CO2 (P.T.N) /ml sangue a 37 ºC.
760
A quantidade de CO2 eliminada a partir do CO2 dissolvido é mínima

e seria incompatível com a vida, haver unicamente transporte de CO2 nesta
forma. O CO2 dissolvido representa cerca de 1/20 do CO2 total presente no
sangue, encontrando-se a maior parte combinado com hemoglobina e sobre- _____
tudo na forma de bicarbonato e ácido carbónico. 19
O azoto não participa nas combustões celulares. Encontra-se unica-
mente dissolvido no organismo com uma pressão parcial igual à do N2 do ar
alveolar, ou seja, PN2 ~- 573 mm Hg. Para 37 ºC o D37 = 0,0132, logo
537
V'' = 0,0132 = 0,01 ml N2 (P.T.N) /ml de sangue a 37 ºC.
760
II.14 - Forças e pressões actuando no tórax e pulmões
Os movimentos do tórax durante o ciclo respiratório são contrariados

por forças de dois tipos distintos, que têm de ser vencidos pela acção dos
músculos respiratórios. Estas forças dividem-se em elásticas e não elásticas
ou viscosas.
Forças elásticas - A Fig. II.7 é, em representação esquemática, um

modelo mecânico do sistema de forças elásticas do pulmão.
Fig. II.7 - Modelo mecânico da respiração.
PT é a pressão no espaço interpleural ou pressão intratorácica. É a

pressão existente no espaço limitado pela pleura visceral e pleura parietal.
A força elástica de retracção dos pulmões expressa por unidade de
_____
superfície de pleura visceral é a pressão pulmonar PL. Na figura anterior

esta pressão é produzida pela mola 1 na base rígida do fole que representa os
pulmões.
A pressão da parede do tórax PW é devida à elasticidade da parede
torácica, com ou sem forças adicionadas aplicadas pelos músculos respira-
tórios. A acção elástica da parede do tórax é simulada pela mola 2.
A pressão no exterior do tórax, representado pelo êmbolo, é a pressão
atmosférica PB e, quando a via da entrada de ar está aberta e não há fluxo de
ar a entrar ou a sair dos pulmões, a pressão interior é também PB.
Quando um orifício é feito na parede do tórax, pondo em contacto
com a pressão atmosférico o espaço existente entre as membranas pleurais,

como ocorre em acidentes, os pulmões colapsam e o tórax expande-se,
devido à entrada de ar naquele espaço. Estes movimentos são devidos à
retracção elástica dos pulmões e à expansão elástica da parede do tórax e
justificam o sentido de actuação das molas 1 e 2, na Fig. II.7 (para o interior
no primeiro caso e para o exterior, no segundo).
Numa representação esquemática mais próxima do real, as pressões
referidas encontram-se indicadas na figura a seguir
Fig. II.8 - Pressões pulmonares.
No fim de uma expiração lenta com a glote aberta, os músculos respi-

ratórios relaxados, as forças elásticas dos pulmões e tórax compensam-se
aproximadamente; os pulmões contêm então cerca de 2 litros de ar.
As forças elásticas dos pulmões, sem oposição, esvaziam os pulmões;
as forças elásticas do tórax expandem-nos até corresponder a um volume
pulmonar de aproximadamente 4 litros. Para volumes pulmonares maiores
do que este, as forças elásticas do tórax actuam para dentro, até à posição
neutra de 4 l.
Opondo-se mutuamente, os pulmões e o tórax formam um sistema elástico,
no qual o trabalho tem que ser produzido pelos músculos respiratórios. _____
Quando não há deslocamento de ar e a via de entrada de ar está
aberta, as pressões equilibram-se, pelo que temos
PL + PT + PB = PW + PT + PB II.49
ou seja,
PL = PW II.50
Se considerarmos unicamente a pleura visceral sob as mesmas

condições, também haverá equilíbrio e podemos escrever, para dois
diferentes estados de inflação
PL1 + PT1 = PB II.51
PL2 + PT2 = PB II.52
Se PL2 é maior do que PL1, como aconteceria para uma maior

inflação pulmonar, PT2 deverá ser menor do que PT1. Por outras palavras, a
pressão intratorácica decresce durante a inspiração. Além disso, como PL
nunca é nula, PT é sempre menor do que a pressão atmosférica PB, na
condição estática com a via de entrada aberta.
Geralmente, PT é medida em relação a PB; PT varia, durante a respi-
ração calma, de 2 cm H2O abaixo de PB até cerca de 7 cm H2O abaixo de
PB, na expiração. Este modo de exprimir a pressão intratorácica PT, leva a
atribuírem-se a esta, valores negativos (-2 e -7 cm H2O, no caso exposto).
Sob certas condições, PT pode ser superior a PB. Durante a expiração
forçada, PT pode ser superior a PB quando o ar é expelido com alta veloci-
dade. Isto consegue-se por contracção dos músculos expiratórios, de modo a
que PW actue fortemente de fora para dentro. Mesmo na condição estática
PT pode ser elevado até valores superiores a PB com um esforço de
expiração contra a glote fechada.
Um violento ataque de tosse, por exemplo, pode elevar PT a 200
mmHg (272 cm H2O) acima de PB.
_____ II.14.1 - Forças elásticas

20
Consideremos um balão feito à custa de um tecido biológico elástico;
bexiga, pulmão lavado ou intestino e debrucemo-nos sobre o seu diagrama
pressão-volume.
Em abcissas está representada a pressão deformadora, ou seja, a pres-
são no interior em relação à pressão atmosférica, e em ordenadas o aumento
de volume, a partir de um volume inicial suposto nulo para uma pressão
deformadora nula.
Todos aqueles materiais apresentam uma curva em S (sigmóide), mais
ou menos pronunciada, como se observa na Fig.II.9. Estas curvas para os

órgãos destacados reflectem as propriedades dos materiais mas podem não
traduzir o seu comportamento no vivo.
Na curva da Fig. II.9, no ponto A, podemos definir as complacências
estática e dinâmica que se aplicam às curvas V(P) não lineares.
Fig. II.9 - Gráfico V(P) para um balão de membrana biológica.
A complacência estática é o coeficiente angular da recta que une a

origem ao ponto A.
AC
Ce = OC II.53
A complacência dinâmica é o coeficiente angular da tangente à curva

V(P) no ponto A
AC
Cd = BC II.54 _____
20
Estas complacências variam ao longo da curva, contudo se a pressão

não ultrapassar E podemos definir complacência linear
DE
C= II.55
OE
constante no intervalo de pressão de O a E.

Os segmentos indicados devem ser interpretados pelos seus valores na

escala dos eixos coordenados, correspondentes às respectivas grandezas, ou
seja, volume e pressão.
Na Fig. II.10, podemos observar a curva V(P) para os pulmões no
vivo, obtida em condições tais que se aproxima de uma recta.
A pressão que tende a encher os pulmões (pressão transpulmonar), ou
seja, a pressão que se opõe a PL, devido à reacção elástica do pulmão, tem o
_
valor PB PT no equilíbrio.
Fig. II.10 - Gráfico volume-pressão para os pulmões (forças elásticas).
A relação entre volume pulmonar e pressão transpulmonar dá informação

sobre a deformação elástica do pulmão. O volume pulmonar é medido com um
espirómetro e PT pode ser determinado por métodos indirectos.
Cada ponto no gráfico, representa um par de medidas, feito sem mo-
vimento do tórax e com a entrada de ar aberta. Com certa aproximação, os
_____
20
pontos encontrados situam-se numa recta.
Chama-se resistência elástica de um pulmão à relação 'P/'V entre a
pressão transpulmonar e a variação do volume pulmonar. Evitaremos esta
designação e usaremos antes a sua relação inversa, ou seja, a complacência
linear do pulmão, cuja definição já vimos.
O trabalho realizado pelas forças elásticas do pulmão será dado pela
soma dos produtos P'V e é igual à área VoBV1.
Recordemos que a área entre a curva e as abcissas no gráfico P(V)
tem as dimensões de uma energia.

De modo análogo, é possível determinar o trabalho das forças elás-
ticas do tórax. Num modelo experimental de sistema respiratório, próximo
de uma situação real, coloca-se o tronco dentro de uma caixa hermética,
onde se varia a pressão, de modo a serem obtidos diferentes graus de insu-
flacção. Os músculos respiratórios são mantidos completamente relaxados.
Este sistema substitui o trabalho dos músculos torácicos.
À diferença de pressão entre o interior da caixa e a atmosfera chama-
se pressão transtorácica. Esta é a pressão que está efectivamente a produzir a
_
distenção do tórax e pulmões durante a respiração, ou seja (PB PC) na Fig.
II.11. Nesta figura esquematiza-se o dispositivo referido.
Fig. II.11 - PC - Pressão no interior da caixa metálica obtida à custa do

êmbolo. PT - Pressão intratorácica. PB - Pressão atmosférica.
_____
Os volumes armazenados nos dois elementos, interior e exterior, são iguais. 20
A diferença de pressão através do sistema total para um dado volume

de gás armazenado é a soma dos dois termos de pressão, ou seja,
PB - PT + PT - PC = PB – PC II.56
O sistema total constituído pelos dois elementos elásticos, pulmão e

parede do tórax, armazena um volume igual aos considerados para as curvas
1 e 2 logo, a curva total 3 pode ser obtida ponto por ponto, através da soma
das pressões das curvas 1 e 2 para cada valor de V.
A área compreendida entre a curva 2 e as abcissas corresponde ao tra-
balho das forças elásticas do tórax.
Das curvas da Fig. II.12 podem calcular-se as complacências dos pul-
mões e caixa torácica, bem como a complacência resultante.
Fig. II.12 - Curva 1 - Característica P(V) do elemento elástico interior, ou se-

ja, o pulmão. Curva 2 - Característica P(V) do elemento elástico
exterior, ou seja, a parede do tórax. Curva 3 - Curva P(V) total.
Dividindo a última expressão pelo valor de V correspondente, vem
_ _ _
PB PT PT PC PB PC
+ = II.57
V V V
_____ ou
20
l l l
+ = II.58
CL CC Cr
onde CL, CC e Cr são, respectivamente, as complacências dos pulmões,

caixa torácica e resultante.
Como se vê, a associação em série das complacências (ou capa-
cidades) dos pulmões e tórax tem como resultante a complacência resultante
Cr, cujo valor se determina de modo idêntico ao usado para o cálculo da
capacidade resultante de uma associação em série de condensadores.

Neste estudo considerámos o volume nulo para uma diferença de
pressão nula, o que não corresponde à realidade. O estudo da situação real é
porém feito em moldes idênticos.
II.14.2 - Forças não elásticas
A respiração envolve forças não elásticas ao movimentar o ar e as

pleuras. Logo, no caso dinâmico, o gráfico do volume do pulmão em função
da pressão deve ser diferente do caso estático, uma vez que neste não se en-
tra com a fricção. O diagrama dinâmico da pressão u volume para um ciclo
completo está representado na Fig. II.13, em que a abcissa é a pressão intra-
torácica, que é igual à pressão transpulmonar multiplicada por -1 depois de
subtraída de PB. Neste gráfico, os pontos B e Vo coincidem com a curva
anterior, pois representam dois pontos estáticos.
Fig. II.13 - Gráfico volume - pressão considerando as forças de fricção.
A inspiração é representada pela linha VoAB e a expiração por _____

BCC’Vo. Nos inícios da inspiração e expiração, há uma grande variação de 20
pressão para uma pequena variação de volume, pois tem que se estabelecer
uma diferença de pressão (da ordem dos 15 cm de água) entre os alvéolos e
a boca, para que o ar se movimente (pressão de abertura).
O trabalho contra as forças de fricção e para movimentar o ar é dado
pelas áreas 2+3 nas Figuras II.14 e II.15. O trabalho total na inspiração (dis-
sipativo e elástico) é dado pela área (1+2+3,) Fig. II.14, e terá de ser
fornecido pelos músculos inspiratórios.
Quando os pulmões atingem o volume Vl, as pleuras estão tensas,
possuindo a energia elástica (Fig. II.14), armazenada nos tecidos. Na expi-
ração, parte desta energia é utilizada para vencer as forças de atrito ao mo-
vimentar o ar e as resistências dos tecidos. A energia necessária para retrair
a parede do tórax é (4). A energia (5) tem de ser fornecida pelos músculos
expiratórios.
Fig. II.14 - Inspiração - Gráfico V(P) considerando as forças de fricção.
O diagrama da Fig. II.15 depende do modo como se faz a expiração

entre os mesmos volumes extremos dos pulmões, Vo e V1. Para uma expira-
ção muito lenta, o trabalho das forças de atrito do ar é muito pequeno,
tornando-se o gráfico muito mais esguio e próximo da linha a tracejado que
separa as áreas 2 e 3. Neste caso, a energia total despendida na expiração é
menor.
_____
20
Fig. II.15 - Expiração - Gráfico V(P) considerando as forças de fricção.
Contudo a fracção da energia elástica armazenada usada de maneira

útil é maior, podendo ser suficiente para realizar a expiração sem a necessi-
dade de trabalho muscular.
Inspiração
1 - Trabalho realizado contra as forças elásticas do tecido.
2 - Trabalho realizado para movimentar o ar.
3 - Trabalho libertado nas resistências viscosas dos tecidos.
Expiração
4 - Trabalho realizado para retrair a parede do tórax.
5 - Trabalho fornecido pelos músculos expiratórios.
Distribuição do trabalho durante a respiração numa pessoa normal:

60-70% - vencer as forças elásticas.
25% - movimentar o ar e vencer as forças de fricção nos tecidos.
II.14.3 - Frequência respiratória e energia dispendida na respiração
Tem interesse saber qual a frequência da respiração que necessita de

menor energia para movimentar uma determinada quantidade de ar por mi-
nuto. O ar expirado por minuto, também chamado ventilação pulmonar,
•
representa-se por VE e é definido para as condições P.T.N., (Fig. II.16). O
•
ponto situado acima do V em VE significa que se trata de uma quantidade
por unidade de tempo.
_____
20
Fig. II.16 - Pressões pulmonares a actuar na expiração. PM é a pressão

da acção muscular (ou a pressão resultante da energia
elástica armazenada) e PA a pressão alveolar.
•
A ventilação pulmonar VE é igual ao volume corrente VT a multi-
plicar pela frequência respiratória, esta, o número de ciclos respiratórios por

unidade de tempo. Na Fig. II.17, o volume corrente é V T = V 1- V o e aparece
em função da pressão pulmonar, ou transpulmonar, que é a pressão a actuar
através da parede pulmonar.
Fig. II.17 - Volume pulmonar em função da pressão.
O trabalho total da respiração pode ser decomposto em trabalho das

forças elásticas e trabalho de forças de fricção.
O trabalho das forças elásticas, por ciclo, é dado pela área do
triângulo VoBV1Vo. A área do triângulo fica 4 vezes maior se VT duplicar,
fica 9 vezes maior se VT triplicar, etc, pois a base e a altura do triângulo
aumentam na mesma proporção. Esta proporcionalidade pode ser
representada por
WEL D (VT)2 II.59
Vimos que:
•
VE = f V T II.60
ou
_____
l
2 VT D f II.61
(O sinal D entre duas quantidades significa proporcionalidade.)
Então, para uma ventilação pulmonar constante, o volume corrente

VT é inversamente proporcional à frequência. Para uma ventilação cons-
tante, se a frequência duplicar, o volume movimentado por ciclo diminui
para metade.
O trabalho realizado por unidade de tempo é, a partir da Eq. II.59.
•
WEL= WEL u f II.62
= (VT)2 u f II.63
l
D ( )2 u f II.64
f
l
D f II.65
Logo, o trabalho das forças elásticas por segundo diminui quando a

frequência aumenta.
O trabalho das forças de fricção por segundo é calculado pela diferen-
ça de pressão, a multiplicar pelo fluxo médio, ou seja,
• •
WFR = (PB – PT) VE II.66
Para o regime laminar (ver Cap. III), que supomos prevalecer na

respiração calma, a pressão motora é proporcional ao caudal
•
P2 - P1 D V II.67
Donde
• • •
WFR = (PB - PT) VE D ( VE)2 II.68
que é independente da frequência. No entanto, o problema fisiológico é mais

•
complicado, atendendo a que interessa uma ventilação alveolar VA cons-
•
tante e não VE = Cte. • •
A ventilação pulmonar VE é igual à soma da ventilação alveolar VA
_____
com a ventilação não efectiva, volume das passagens de ar VD, multiplicado
2
pela frequência respiratória.
Assim, o trabalho das forças de fricção por unidade de tempo vem
• • • •
WFR D ( VE)2 ou WFR D (VA+ VDf )2
ou ainda
• • • •
WFR D (VA)2 + 2VAVDf + (VE)2 f 2 II.69
que indica que a energia dissipada por segundo, na forma de calor, aumenta
com a frequência.
O valor mínimo do trabalho total por unidade de tempo coincide com
a frequência respiratória normal, quer no repouso (ventilação de 12 "/min),
quer no exercício moderado (ventilação de 25 "/min). Verifica-se, portanto,
que as frequências respiratórias adoptadas naturalmente para as diferentes
necessidades ventilatórias correspondem ao máximo rendimento energético.
Na prática, não é fácil verificar que o trabalho das forças elásticas é
l •
exactamente proporcional a , pois quando f cresce, VE cresce também. No
f
entanto, como indica a Fig. II.18, existe uma frequência para cada ventilação
pulmonar a que corresponde um trabalho mínimo.
Fig. II.18 - Trabalho respiratório em função da frequência.
_____
21
II.15 - Física do alvéolo-pulmonar
Consideremos um pulmão destacado da caixa torácica e estudemos a

curva V(P), supondo que as medidas são feitas sem movimentos bruscos e
sem obstáculos à entrada de ar no pulmão. Nestas condições, a característica
V(P) apresenta um ciclo de histerese, ou seja, a curva de preenchimento é
diferente da curva de esvaziamento. Isto não é previsível pois, não obstante
a complexidade do sistema, tudo levaria a pensar que as curvas V(P) fossem
coincidentes nos dois sentidos, havendo o cuidado de fazer as medidas em

condições de quase repouso, para as quais a resistência do ar e outras fric-
ções não seriam factores de grande importância. Por outras palavras, criando
sucessivas condições de quase-equilíbrio, a que correspondem valores de P
e V, durante o enchimento, não parece lógico que estes pares sejam diferen-
tes, para os mesmos valores de V, durante o esvaziamento.
A parte do sistema respiratório em estudo é constituída por um tubo
grosso, a traqueia, que se ramifica dando os brônquios, estes por sua vez os
bronquíolos que, numa longa sequência de divisões dão origem aos tubos
alveolares e alvéolos. Os alvéolos são portanto os receptores finais do ar inalado
abrindo-se e dilatando quando recebem o ar e diminuindo de volume quando o
expiram. Podemos pois dizer que os alvéolos pulmonares, pequenas bolsas
aproximadamente esféricas, onde o ar e o sangue estão separados por uma fina
parede, são dilatados pelo ar durante a inspiração e esvaziam quase
completamente durante a expiração. Num pulmão os alvéolos têm diversos
volumes, havendo diferenças consideráveis nos seus raios.
As paredes do alvéolo são constituídas por uma membrana muito fina
sobre a qual se encontra uma camada líquida.
Fig. II.19 - Esquema da constituição de um alvéolo pulmonar.
_____
O comportamento mecânico do alvéolo não depende exclusivamente 21
da elastância da membrana pulmonar, pois é resultante das propriedades do
tecido e da superfície líquida, actuando em simultâneo. Este facto foi
demonstrado pela primeira vez por Neergard em 1928, que verificou que as
curvas V(P) para o pulmão destacado eram muito diferentes quando preen-
chia o volume do pulmão com ar ou soro fisiológico, Fig. II.20.
Pelo facto do raio alveolar ser muito pequeno e a membrana ser muito
fina, são as propriedades superficiais da camada líquida (ou seja, a tensão
superficial do líquido) que têm maior preponderância no processo, podendo
dizer-se que a sua contribuição é da ordem dos 3/4. É a tensão superficial da

camada líquida que explica, não só a curva de histerese anteriormente refe-
rida, como também o facto de o ar não passar dos alvéolos de menor raio
para os de maior raio, como seria de prever pela fórmula de Laplace3.
Fig. II.20 - Curvas V(P) para o pulmão destacado com volume preenchido
com ar e soro fisiológico (Neergard).
Como veremos, o papel dos fenómenos de superfície no comportamento

do alvéolo é vital. De um modo geral, podemos dizer que as propriedades da
camada líquida superficial do alvéolo impedem o colapsamento, permitem a
coexistência de alvéolos de raios diferentes, facilitam a difusão do oxigénio,
aumentam a complacência pulmonar, diminuem o trabalho respiratório e
contribuem para a estabilização das trocas de fluidos no alvéolo.
Vejamos como se comportam individualmente as duas camadas do alvéolo.
Quanto à membrana, a variação do volume em função da pressão
transalveolar para o epitélio de um alvéolo com raio ro e volume Vo (para
uma pressão mínima sem colapsamento), de acordo com o que vimos
(II.14.1), ocorre com uma variação aproximadamente linear até ser atingida
uma região onde a deformação passa a ser mais lenta (Fig. II.21).
Sobre a camada líquida, principal responsável pelas propriedades do
_____ alvéolo, há bastante para dizer. De facto, as propriedades de superfície do
21 líquido que reveste o alvéolo são importantes para a compreensão dos resul-
tados experimentais, pois a tensão superficial desta solução não é constante
mas varia quando varia a área do alvéolo.
É sabido que, para certas soluções, existe a tendência para haver uma
maior concentração de moléculas de soluto à superfície e próximo da
superfície. Isto resulta do facto das interacções que as moléculas do soluto
exercem sobre as do solvente serem menos intensas do que as interacções
3
- Neste estudo interessa o conhecimento das propriedades das superfícies líquidas que são
tratadas em detalhe no Cap. IV.
entre as próprias moléculas do solvente. Do balanço entre este fenómeno e a

difusão molecular resulta uma diminuição da tensão superficial do líquido.
Fig. II.21 - Variação do volume em função da pressão transalveolar para o epitélio de um

alvéolo com volume Vo para uma pressão mínima sem colapsamento.
Resulta também que a concentração do soluto à superfície do líquido

é maior do que no interior. Neste tipo de moléculas, se a área da superfície
do líquido aumentar, ocorre uma redistribuição, de modo que a concentração
à superfície é reconstituída à custa de moléculas mais interiores de soluto,
não variando a tensão superficial da solução.
Existe um outro tipo de moléculas, com propriedades ditas de
surfatante (em inglês “surfactant”, contracção de “surface active agent”) que
em solução, além de baixarem a tensão superficial, originam que esta varie
com a área da superfície livre do líquido.
_____
21
Fig. II.22 – di-palmitoíl-fosfatidil-colina.

Estas substâncias são fosfolípidos que têm conformação longitudinal

e apresentam, num dos seus extremos, uma região hidrofílica e no outro
extremo uma região hidrofóbica apresentando uma notável tendência para se
situarem à superfície da água. Na Fig. II.22 encontra-se representada uma
molécula de um fosfolípido, a di-palmitoíl-fosfatidil-colina.
Após a dissolução de uma pequena massa de um fosfolípido em água
ocorre ligação de moléculas deste com as moléculas de água à superfície,
feita à custa dos grupos hidrofílicos (ácidos gordos), ficando os grupos hi-
drofóbicos (colina) ''fora da solução'' virados para o exterior. Dado que estas
ligações são de pequena energia as moléculas do surfatante ficam em grande
quantidade à superfície originando uma considerável diminuição da tensão
superficial. No interior do líquido estas moléculas aglomeram-se formando
partículas onde as moléculas de água não penetram, pois os grupos
hidrofóbicos viram-se todos para o interior, ficando os hidrofílicos virados
para o exterior e ligados a moléculas de água, Fig. II.23. Formam-se também
lipossomas, de dupla camada, com solvente no interior.
Fig. II.23 - Após a dissolução de uma pequena massa de um fosfolípido em água,

ocorre ligação de moléculas deste com as moléculas de água à superfície,
feita à custa dos grupos hidrofílicos. No interior do líquido estas molécu-
las aglomeram-se, formando partículas, limitadas por uma monocamada,
_____ onde as moléculas de água não penetram e lipossomas, de dupla camada,
21 com solvente no interior.
As partículas formadas no interior do líquido, são insolúveis e cons-

tituem associações moleculares com propriedades semelhantes às de fibras
elásticas. No caso da área da superfície aumentar, estas partículas não podem
ceder moléculas para reconstituir a concentração à superfície. Assim, quando a
área da superfície livre do líquido aumenta, por exemplo devido a uma acção
mecânica, as moléculas do soluto são substituídas por moléculas de água, a
concentração do soluto à superfície diminui e aumenta a tensão superficial da
solução (Fig. II.24).
No alvéolo as "células alveolares do tipo II", que ficam no interior dos

alvéolos, produzem uma mistura de fosfolipídos (~90%) e proteínas (~10%)
que constituem o surfatante pulmonar.
Fig. II.24 - Quando se reduz a área da superfície livre de uma solução de surfatante, a
concentração do soluto à superfície diminui e aumenta a tensão superficial
da solução pois, não havendo moléculas de soluto disponíveis, são
moléculas de água que se dirigem para a superfície. Na figura, as molé-
culas de água (solvente) estão representadas por pequenos círculos negros.
Nas moléculas de surfatante os braços representam a parte hidrofóbica e
as pequenas circunferências a parte hidrofílica.
Notemos que a variação da tensão à superficie resultante da variação

da área superficial é uma propriedade que associámos às membranas
elásticas e não às superfícies líquidas.
Se representarmos a variação da tensão numa membrana elástica, com
um único tipo de fibras elásticas, em função da área da membrana (Fig.
II.25), temos uma recta com uma inclinação, que depende do coeficiente de
elasticidade do material. Para um líquido puro a tensão superficial não
depende da área.
_____
21
Fig. II.25 - Comparação da variação da tensão superficial de um líquido e da

tensão elástica de uma membrana com a superfície.
Se fizermos a representação T(S) para uma solução de surfatante (ou

para o líquido que cobre os alvéolos), a curva que se obtém apresenta um
ciclo de histerese, Fig. II.26.
Quando a área aumenta, ou seja, se passa de A para B pelo trajecto 1,
a tensão superficial aumenta rapidamente até ser atingido o valor constante
Vs que corresponde à tensão superficial do solvente. A tensão superficial da
camada líquida do alvéolo pulmonar varia de cerca de 5 dine/cm, quando a
área do alvéolo é mínima, até cerca de 50 dine/cm quando esta é máxima.
Quando ocorre diminuição da área, trajecto de B para A (percurso 2),
a inclinação da curva é menor do que na parte ascendente de 1 e não existe
uma região paralela às abcissas como no caso anterior.
Fig. II.26 - Ciclo de histerese da tensão superficial do líquido alveolar em

função da área interior do alvéolo.
A explicação da curva V(S), apresentada por soluções de surfatantes,

parece estar não só no comportamento da superfície, como nas associações
moleculares criadas no interior, com propriedades elásticas, cujos efeitos
conjugados resultariam em propriedades intermediárias entre membranas e
superfícies líquidas.
_____ Consideremos a curva V(P) para uma película de um líquido sem
21 surfatante (solução de sabão, por exemplo), colocada na extremidade de um
tubo cilíndrico. Esta curva toma o aspecto da Fig. II.27 a), com explicação
na Fig. II.27 b). Considera-se que V é o volume limitado pelo interior da
película, suposta uma calote esférica de raio r, e o plano tangente à entrada do
tubo. A pressão P é obtida aplicando a Eq. IV.11, entrando com o raio da
calote. Para pequenos volumes (curvas 1 e 2) os raios de curvatura da calote
são grandes e as pressões pequenas. A pressão só pode aumentar até um
valor máxima que corresponde ao valor mínimo do raio da calote, ou seja,
quando é igualado o raio do tubo (curva 3). A partir daqui a pressão volta a
diminuir com aumento do volume (curvas 4 e 5).
A curva da Fig. II.27 a) depende da tensão superficial da solução. Para
bolas de sabão com tensão superficial numa sequência de valores crescentes
(numeradas de 1 a 5), as curvas V(P) seriam do tipo da Fig. II. 27 c). Para os
mesmos valores de V, os valores de P aumentam com o aumento da tensão
superficial (valores crescente de 1 a 5 podendo 5 corresponder ao solvente).
Fig. II.27 – a) Curva V(P) para uma bola de sabão. b) Detalhes: 1–Volume mínimo; Pequena
pressão; Grande raio de curvatura do alvéolo. 2 – Aumento da pressão; Maior
volume; Menor raio. 3 - Raio mínimo (iguala o raio do tubo); Pressão máxima.
4 e 5 - O volume e o raio de curvatura aumentam e a pressão diminui. c) Curvas
V(P) para bolas de sabão com tensões superficiais com valores crescentes
(curvas de 1 para 5).
Para uma solução de surfatante a curva V(P) numa experiência

idêntica à da Fig. II.27 b) seria diferente. Neste caso, com o aumento de V a
tensão superficial da solução iria aumentar, (Fig. II.28 a)), tendendo para o _____
21
valor máximo Vs, tensão superficial do solvente. Nesta curva onde o volume
limitado pela película é definido como no caso anterior, para um tubo de raio
r, observa-se uma marcada histerese. Na Fig. II.28 b) encontra-se um conjunto
de curvas V(P), para valores de V a variar desde um valor mínimo V1 até Vs,
tensão superficial do solvente. Os valores da pressão para qualquer valor do
volume limitado pela membrana da solução de surfatante podem ser obtidos a
partir das curvas da Fig. II.28 b) desde que se escolha a curva que corresponde
ao correcto valor da tensão superficial. Por exemplo para o valor de volume
V10 (Fig. II.28 a)), do esvasiamento, a que corresponde o valor de tensão

superficial V10 o valor de pressão no interior é P10 (Fig. II.28 b)).
Fig. II.28 – a) Curva V V) para solução de surfatante. b) Curvas V(P) paraFDPDGDV

líquidas de tensão superficial constante e de valores V1, V2,..Vi,.. Vs. c)
Curva V(P) para uma camada líquida de solução de surfatante (cheio grosso).
Para o volume V10 no esvasiamento, a que corresponde o valor de tensão
superficial V10 (a)), o valor de pressão no interior é P10, (b)). O par
(V10,P10) é o ponto 10 da curva V(P) de esvasiamento para a película de
VROXomRGHVXUIDWDQWH F
Procedendo de igual modo para um conjunto de pontos obtêm-se

pares de valores (Pi,Vi) que correspondem à curva V(P) (Fig. II.28 c)) para
uma película de solução de surfatante com a histerese que seria de prever.
A curva da Fig. II.28 c) simula o comportamento da camada líquida do
alvéolo no ciclo respiratório. Durante a inspiração, o seu volume aumenta,
aumentando a tensão superficial de acordo com a Fig. II.28 a), e o inverso na
_____
expiração.
2 O comportamento do alvéolo depende, como vimos, da soma das
propriedades das duas camadas - a membrana alveolar e a película líquida,
cujo estudo acabámos de fazer.
Consideremos a contribuição da membrana elástica.
Na Fig. II.29 apresentamos as curvas V(P) para uma membrana
elástica com o nº 2 e para uma camada líquida sem surfatante com o nº 1,
como se indicou na Fig. II.19.
Para cada valor do volume, a pressão do sistema membrana-camada
líquida é a soma das pressões nas curvas nºs 1 e 2. A curva nº 3 é a curva
soma e representa a função V(P) para o sistema constituído por uma
membrana elástica coberta com uma camada líquida de uma solução sem
surfatante4. Notar que nesta curva ocorre um ligeiro efeito de histerese.
Considerando o enchimento por aumento contínuo da pressão, o trajecto a
seria o único que poderia ocorrer. No esvaziamento por decréscimo contínuo
da pressão ocorreria o trajecto b.
Fig. II.29 - Curvas V(P) para: 1 - Camada líquida sem surfatante; 2 - membrana
alveolar sem qualquer líquido; 3 - soma 1+2.
Consideremos, finalmente, a situação de uma membrana elástica com

uma camada líquida aderente de uma solução de surfatante.
_____
2
Fig. II.30 - Curvas V(P) para: 1 - Camada líquida sem surfatante; 2 membrana
alveolar sem líquido; 3 - soma 1+2.
4
- Supõe-se desprezível a acção da tensão de contacto líquido-membrana.
Na Fig. II.30 encontram-se as curvas V(P) para a camada líquida com

o seu ciclo de histerese (Curva 1, igual à da Fig. II.29 c)), para a membrana
(Curva 2) e para o sistema membrana camada líquida, ou seja, a curva soma
(Curva 3). A curva soma obtém-se considerando em separado as duas
componentes (ascendente e descendente) da curva 1. De realçar o marcado
efeito de histerese.
A curva 3 (Fig. II.30) aproxima-se da curva de enchimento do alvéolo.
Uma observação importante quando se comparam as curvas V(P) com
líquido sem surfatante (Fig. II.29 curva 3) e V(P) com solução de surfatante
(Fig. II.30 curva 3) é que, na primeira, a pressão de colapsamento do alvéolo
é consideravelmente superior à obtida com surfatante.
Durante a expiração são atingidas pressões nos alvéolos que levariam
ao total colapsamento se não existisse surfatante.
Quando se considera a curva V(P), agora para os pulmões no seu
total, observa-se igualmente um pronunciado efeito de histerese, Fig. II.31.
O volume pulmonar para dada pressão é nitidamente mais elevado durante a
expiração do que durante a inspiração.
Fig.II.31 – Curvas V(P) para o enchimento e esvaziamento do pulmão.
_____
Além do que já estudamos sobre o alvéolo, um outro facto contribui
22
para a diferença entre as curvas V(P) no enchimento e esvaziamento do
pulmão. Como os raios dos bronquíolos são diferentes, as pressões de
abertura são diferentes de alvéolo para alvéolo. Ao aumentar-se a pressão há
um recrutamento progressivo dos alvéolos, cujos bronquíolos tenham raios
progressivamente menores. Só depois de atingida a pressão de abertura é
que se processa um aumento rápido do volume para cada alvéolo.
Não existe fenómeno equivalente a este recrutamento durante o esva-
ziamento. Quando se inicia a expiração os alvéolos encontram-se preenchi-
dos e todos à mesma pressão, havendo sincronismo na variação de volume

quando baixa a pressão.
Fig. II.32 – Curvas V(P) para o pulmão com ênfase para

o comportamento nas pequenas pressões.
Os volumes próximos de zero, a que corresponde o colapsamento,

ocorrem para valores de pressão tão baixos, que não são nunca atingidos no
fim da expiração. Se o final do esvaziamento corresponder ao ponto 1, o
enchimento irá processar-se a partir do ponto 2, (Fig. II.32).
É de realçar a eficiência desta camada líquida que desempenha uma
tarefa vital sem protecções especiais e tendo como participação biológica
única, a produção de surfatante pelas células epiteliais do tipo II.
Na vida fetal ocorre produção de surfatante alveolar a partir do 6º
mês, ou mesmo antes, viabilizando o nascimento de prematuros.
_____
22
Fig. II.33 – Curvas V(P) para o líquido alveolar de um

recém-nascido com doença da membrana hialina.
No nascimento os alvéolos estão praticamente colapsados e o seu pri-

meiro enchimento requer um esforço considerável. Uma baixa concentração

de surfatante no recém nascido, como ocorre na chamada doença da
membrana hialina pode ser fatal. Na Fig. II.33 encontra-se a curva tensão
superficial em função da área, V(S), para o líquido alveolar de um recém
nascido com a referida doença. Compare-se com a curva da Fig. II.26.
.
II.16 - Capacidade de difusão de um gás
Numa fase gasosa a velocidade de difusão de uma dada espécie mo-

lecular é inversamente proporcional à raiz quadrada da respectiva massa mo-
lecular (ou da sua densidade). Por exemplo, a velocidade de difusão, em fase
gasosa do CO2 é cerca de 1,7 vezes maior do que a do Xe, valor que é a
relação inversa das raízes quadradas dos seus números de massa.
Numa fase líquida a velocidade de difusão (ou corrente de difusão
molecular) de uma dada espécie molecular gasosa é proporcional ao inverso
da raiz quadrada da sua massa molecular e ao coeficiente de solubilidade do
gás no líquido. Combinando duas expressões conhecidas, a da corrente de
difusão molecular livre e monodireccional (1ª lei de Fick), para uma dada
temperatura t,
dC
I = - D S dx II.70
onde I é a corrente de difusão em moles/s através da superfície de difusão S,

D é a constante de difusão, x é a distância na direcção de difusão e C é a
_____ concentração em x, e a da lei de Henry, para a mesma temperatura
22
& OtP II.71
com Ot coeficiente de solubilidade do gás no líquido, vem
dP
I = - D Ot S dx II.72
O coeficiente de solubilidade mais usado é o coeficiente de solubi-

lidade de Ostwald Ot que é o volume de gás, em ml à pressão de 1 atmosfera

e à temperatura t da experiência que, após equilíbrio, se dissolve em 1ml do
solvente, quando a pressão parcial do gás é 1 atm.
A constante de difusão D é proporcional ao inverso da raiz quadrada
da massa molecular, pelo que a velocidade de difusão relativa de dois gases,
na mesma fase líquida (água, por exemplo) poderá obter-se, para iguais gra-
dientes de pressão parcial e iguais áreas de difusão.
I M´ Ot
= II.73
I´ M O t́
O coeficiente de solubilidade de Ostwald a 37 ºC para o caso do O2 é

O37=0,0244 ml STPD/ml H2O e para o Xe, O37 = 0,084 ml STPD/ml H2O.
Então a velocidade de difusão relativa dos dois gases, na mesma fase líquida
(água por exemplo) poderá obter-se, considerando o produto
32 0,084
u = 1,69 II.74
133 0,0244
ou seja, em fase aquosa, o Xe é 1,69 vezes mais difusível do que o O2 na

mesma fase.
Consideremos agora as velocidades de difusão do Xe em fase aquosa e
em matéria gorda. Para esta matéria o coeficiente de solubilidade de Ostwald a
37 ºC para o Xe é O37 = 1,004. Então a relação entre as velocidades de difusão
do gás nos dois meios poderá obter-se, considerando o cociente
1,1004
= 13,1 II.75 _____
0,084
22
ou seja, o Xe é 13,1 vezes mais difusível em fase gorda que em fase aquosa.
Em termos gerais, na patologia intersticial o valor de O37 para o Xe
deve aumentar em relação ao tecido são, devido ao aumento de colagéneo na
BAC, característico nestas afecções.
Em 1909 Bohr introduziu o conceito de capacidade de difusão do
pulmão, ou seja, o número de ml de um gás (STPD) que difunde através da
membrana pulmonar por minuto e por mmHg de diferença de pressão
parcial entre a mistura gasosa alveolar e o sangue capilar.
Chamando DXe à capacidade de difusão para o Xe, temos, por simples

aplicação da definição
•
VXe
DXe = P II.76
AXe - PCXe
•
Onde VXe é a corrente de difusão molecular pulmonar total de Xe, PAXe é a
pressão parcial do Xe alveolar e PCXe a pressão parcial média do Xe nos
capilares pulmonares.
Por outro lado, numa perspectiva morfométrica, pode escrever-se
Ot S
DXe = Ko II.77
M 'x
com Ko constante. A partir das Eqs. II.76 e II.77 verifica-se que, se conhecendo
•
qualquer dos pares VXe e (PAXe - PCXe) ou S e 'x, pode calcular-se DXe.
II.17 - Difusão através da parede de um capilar pulmonar
Em cada ciclo cardíaco a difusão do O2 e CO2 através da barreira

alvéolo capilar ocorre em fracções de segundo nos trajectos dos capilares em
contacto com os alvéolos. Estes trajectos, que são da ordem dos décimos do
mm, devem permitir a troca de uma fracção substancial dos gases
dissolvidos no sangue capilar para que o processo seja rentável. É possível
fazer o cálculo teórico da eficiência da troca gasosa utilizando um modelo
_____ geral baseado na Fig. II.34.
22 A parede de um tubo capilar, de raio R, é atravessada por uma
densidade de corrente de um soluto, de JA moles cm-2s-1. A permeabilidade
da parede ao gás é Pac cm s-1 e a sua concentração extravascular é CA moles
•
cm-3. O caudal de sangue através do capilar é Q cm3 s-1.
Seja o eixo do tubo, um eixo orientado (xx), com origem no início do
tubo e consideremos duas secções rectas de abcissas x e x+dx, onde ocorrem
concentrações C(x) e C(x+dx) (Fig. II.34).
Fig. II.34 - Modelo de difusão através da parede de um tubo capilar.
As densidades de corrente através de secções rectas do tubo, J S(x+dx)

e JS(x), em dois pontos muito próximos, são diferentes devido à difusão de
gás através da parede. Utilizando as concentrações do soluto em x+dx e x
pode exprimir-se a diferença entre aquelas densidades de corrente pela
equação:
1
[.
JS(x+dx) - JS(x) = A Q C(x+dx) - Q C(x)
.
]
II.78
ou
.
Q
JS(x+dx) - JS(x) = A dC(x) II.79
A expressão da densidade de corrente através da membrana do

capilar, é
JA = Pac (CA - C(x)) II.80
Então, supondo que só ocorrem trocas por difusão em metade da área

lateral dos capilares, já que estes assentam sobre os alvéolos, é também
[JS(x+dx) - JS(x)]SR2 = [Pac(CA - C(x)) ] Sdx _____

ou 22
JS(x+dx) - JS(x) = Pac(CA - C(x)) dx / R II.81
.
Donde, considerando a Eq. II.79 e recordando que Q = SR2 v- com v- a
velocidade média do líquido no tubo
dC(x) P
= - ac (CA - C(x)) II.82
dx vR
ou
dC(x)
+ k C(x) = k CA II.83
dx
e
P
C(x) = CA (1- e- kx) e, como k = - ac II.84
vR
vem
Pac
C(x) = CA ª¬1 - e- ( v- R x)º¼ II.85
Numa situação hipotética, considerando na equação anterior a veloci-

dade média v- (ou R) como parâmetro, com valores de 1, 3, 5, 7 e 9 unidades
arbitrárias e mantendo constantes e com valores razoáveis Pac e R (ou v) -
obtêm-se as curvas seguintes para a variação relativa da concentração do gás
ao longo do capilar (Fig. II.35).
Fig. II.35 - Curvas C(x)/CA em função do comprimento do capilar, para

-
diversos valores de velocidade média v.
_____
22
Do mesmo modo, considerando como parâmetro a permeabilidade
com valores de 1, 3, 5, 7 e 9 unidades arbitrárias, obtêm-se as curvas seguin-
tes para a variação relativa da concentração em função do comprimento x do
capilar (Fig. II.36).
As curvas C(x)/CA vs. x tendem mais rapidamente para a saturação
com a diminuição da velocidade e com o aumento da permeabilidade da
membrana.
Fig. II.36 - Curvas C(x)/CA em função do comprimento do capilar, para di-

versos valores de permeabilidade Pac.
Consideremos agora a situação humana.

No caso do N2 da mistura gasosa alveolar, Pac = 0,13 cm s-1. Os
valores médios do raio do capilar alveolar e velocidade do sangue são,
respectivamente R = 4u10-4 cm e v- = 0,04 cm s-1. Calculemos o valor da
relação entre as concentrações C(x)/CA ou entre as correspondentes
pressões parciais do gás P(x)/PA , para alguns valores de x.
Para x = 10 Pm P(x)/PA = 0,999703955

Para x = 20 Pm P(x)/PA = 0,999999912
Para x = 50 Pm P(x)/PA = 0,999999999
Para o caso do CO2, Pac = 2,6 cm s-1. Supondo condições

semelhantes ao caso anterior a relação entre as pressões parciais P(x)/PA ,
para x = 10 Pm já é praticamente unitária.
_____
II.18 - Tubo colapsável ideal 22
Um tubo colapsável ideal é uma entidade teórica que possui as se-

guintes propriedades: é altamente deformável, não é elástico, mantém-se
aberto, por completo, para qualquer pressão transmural positiva e colapsa
para qualquer pressão transmural negativa.
Consideremos três situações típicas de funcionamento destes tubos:
1. - Quando a pressão no meio circundante PS de um tubo colapsável
é superior à pressão à entrada do tubo Pi, não ocorre caudal através do tubo,
Fig. II.37, ou seja
PS > Pi > Po
e
.
Q=0 II.86
Fig. II.37 - Comportamento do tubo colapsável ideal para o caso de P S > Pi >
Po. Pi - Pressão à entrada do tubo; Po - Pressão à saída; PS -
Pressão circundante.
2. - Quando, como na Fig. II.38, a pressão à saída do tubo é maior do

que a pressão no meio circundante, o caudal é proporcional à diferença entre
a pressão de entrada e a pressão de saída e variações na pressão exterior,
mantendo-se a condição inicial, não influenciam o caudal. Neste caso
Pi > PS > Po II.87

e o caudal é
.
Q = ( Pi - Po ) / R II.88
com R - Resistência hidrodinâmica.
_____
2
Fig. II.38 - Comportamento do tubo colapsável ideal para o caso de P i > Po >
PS; Pi - Pressão à entrada do tubo; Po - Pressão à saída; PS -
Pressão circundante.
3. - Quando um tubo colapsável é rodeado por uma pressão maior do

que a pressão à saída do tubo, mas menor do que a pressão à entrada, o tubo
vai colapsar na extremidade distal. Uma alteração do caudal não vai influen-
ciar a pressão no extremo colapsado do tubo e uma variação na queda de
pressão através do tubo não influencia o caudal. Neste caso
Pi > PS > Po II.89
e o caudal é dado por
.
Q = ( Pi - PS ) / R II.90
Este comportamento é conhecido por resistência de Starling.

A circulação pulmonar é o exemplo clássico da importância fisioló-
gica dos tubos colapsáveis.
Fig. II.39 - Comportamento do tubo colapsável ideal para o caso de P i > PS >
Po; Pi - Pressão à entrada do tubo; Po - Pressão à saída; PS -
Pressão circundante. R - Resistência hidrodinâmica.
_____
2
Considerando o pulmão dividido em três terços (Fig. II.40). Palv, Part e
Pven representam respectivamente a pressões alveolar, arterial capilar e ve-
nosa capilar. Verifica-se que no terço superior (I) Palv > Part > Pven não ha-
vendo caudal sanguíneo através do capilar. No terço médio (II) Part > Palv >
Pven ocorrendo caudal cujo valor é proporcional a Part - Palv. Nesta região a
pressão Pven não influencia o caudal. É a chamada região da resistência de
Starling. No terço inferior (III) Part > Pven > Palv. O caudal não é influen-
ciado pela pressão Palv.
No indivíduo normal em pé a Zona I não ocorre. Pode, no entanto
observar-se em situações patológicas, quando a pressão arterial pulmonar
baixa ou Palv é elevada.
Fig. II.40 - Consequências do efeito hemostático nos capilares alveo-

lares considerados tubos colapsáveis ideais.
II.19 - Difusão dos gases nas vias respiratórias
Considere-se que na inspiração existe, à entrada, uma variação abru-

pta entre as concentrações do O2 e CO2 no ar que entra e no gás retido nos
pulmões. Por exemplo, para o CO2, 0% no ar inspirado e 5% no gás retido
nos pulmões no fim da última expiração. Na inspiração, nas primeiras gera-
_____
ções das vias de ar e devido à corrente de arrastamento esta frente desloca-se
23
rapidamente à medida que se desfaz e alarga. Devido ao aumento de área a
velocidade diminui progressivamente até ficar com valor mínimo, cerca da
17ª ou 18ª geração. Cria-se aí uma competição entre o caudal de arrasta-
mento, tendendo a mover o gás inspirado para o alvéolo e a difusão dos
gases alveolares, em direcção à boca. No final da inspiração há ainda uma
marcada diferença entre as concentrações dos gases nas vias mais proximais,
até cerca da 15ª geração onde a composição é próxima da do ar ambiente e
as das vias mais periféricas e alvéolos. Esta diferença é conhecida por
estratificação. No início da expiração a estratificação desaparece rapida-

mente: no caso do CO2, difusão e arrastamento contribuem simultaneamente
para desfazer o gradiente de concentração e deslocar a frente de concen-
tração em relação ao exterior.
II.20 - Modelo mecânico avançado do transporte gasoso na respiração
O modelo mecânico da respiração, apresentado na Fig. II.7, só

pretende abarcar os aspectos mais globais da dinâmica dos gases na
respiração. Modelos especializados podem ser mais esclarecedores da
mecânica dos processos envolvidos.
Antes de mais, consideremos o comportamento de um tubo elástico T,
Fig. II.41, situado numa câmara onde existe a pressão Pe, percorrido por um
caudal gasoso F, com regime laminar e determinando a pressão Pi(x) no seu
interior. A pressão do gás à entrada do tubo é Pi(0) e à saída é Pa, a pressão
atmosférica. Vamos supor ainda que podemos variar o valor da pressão Pe
da câmara. Na Fig.II.41 a pressão Pi(0) > Pe, pelo que o tubo inicialmente
dilata, passando o raio a diminuir a partir do valor de x em que Pe > Pi(x).
Na Fig. II.42 encontram-se, a traço cheio fino, as curvas do caudal em
função da queda total de tensão ao longo do tubo [Pib(0) = Pi(0) – Pa], tendo
como parâmetro o valor da pressão barométrica na câmara [Peb = Pe – Pa]. A
inclinação destas curvas, em cada ponto tem as dimensões do inverso de
uma resistência hidrodinâmica. Com o aumento do parâmetro Peb e para o
mesmo Pib(0) o diâmetro do tubo diminui, (a resistência hidrodinâmica
aumenta) pelo que F será menor. Evidentemente, se o tubo fosse rígido,
teríamos uma só curva, uma recta, a passar pela origem.
_____
23

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SRU XP FDXGDO JDVRVR HP UHJLPHODPLQDU )FRPDSUHVVmR3 i [
QRVHXLQWHULRU3apDSUHVVmRDWPRVIpULFD
Se para cada uma das curvas F [Pib(0)] considerarmos os pontos para

os quais Peb = Pib(0) e traçarmos a curva que une estes pontos, obtém-se a
curva a traço cheio médio, da Fig. II.42.
)LJ,,$FKHLRILQRDVFXUYDVGRFDXGDO)YVDTXHGDWRWDOGHWHQVmRDWUDYpV
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Esta curva representa valores de caudal em função de Pib(0), mas com Pe

= Pi(0), ou seja, a curva que seria obtida para estas variáveis, com o sistema da
Fig, II.43, onde o anel perfurado S, suporta a membrana e permite ter Pe = Pi(0).
(VWDFXUYDDSUHVHQWDXPPi[LPRSDUDRFDXGDO)mTXHRFRUUHSDUDR
YDORUGHDEFLVVD3f(VWHPi[LPRUHVXOWDGDGXSODDFomRGRDXPHQWRGHPe =
Pi(0), ou seja, o aumento do caudal e da acção constritora sobre a parede
elástica. Para pressões baixas predomina o primeiro efeito, para pressões
elevadas, o segundo. Estamos a considerar valores positivos de Peb pois os
_____ valores negativos, equivalem ao caudal gasoso em sentido inverso, não
23 ocorrendo extremos na curva, nesta região.
)LJ,,D 6LVWHPDRQGHRDQHOSHUIXUDGR6VXSRUWDDPHPEUDQDHSHUPLWH
WHU3H 3L E &XUYD) 3HE).
Para aproximar mais o modelo da situação real do movimento do gás

nas vias aéreas, interessa adicionar uma resistência hidrodinâmica RH, em
série, que vai representar, sobretudo, o comportamento dos tubos rígidos das
vias aéreas (traqueia e tubos das últimas gerações), Fig. II.44. A curva 2 é a
curva total da série e a curva 3 é da componente resistiva, uma recta como
seria de esperar. A curva 2 é a soma das curvas 1 e 3.
Fig. II.44 - a) Sistema da Fig. II adicionada de um componente resistivo, RH;

b)&XUYD) 3HE) para o sistema (2) e para RH (3). _____
23
Finalmente considere-se o modelo da Fig. II.45. O tubo T que utilizá-

mos na Fig. II.44, encontra-se agora colocado numa câmara estanque
elástica E, de volume V. O tubo e a câmara estão ligados de forma elástica,
sendo esta ligação representada por molas fixadas nestes elementos.
Fig. II.45 - O tubo T da Fig. II.44 encontra-se colocado numa câmara estan-
que elástica E de volume V. O tubo e a câmara estão ligados de
forma elástica sendo esta ligação representada por molas fixadas
nestes elementos.
Quando o êmbolo B é deslocado, varia a pressão Ppl e também a pres-

são Pi, por deformação de E. Se Ppl > Pi a câmara E colapsa, ocorrendo dila-
tação na situação oposta. A pressão Pi é transmitida ao início do tubo T ori-
ginando caudal gasoso que depende de Ppl, e pode ser nos dois sentidos. A
pressão a actuar efectivamente na parede E é Ppl – Pi, que pode ser positiva
ou negativa. Esta diferença de pressão está relacionada com o volume V da
câmara E de forma linear, pela equação
Ppl – Pi = a(V – Vo) II.91
onde a é uma constante que depende das características de E e Vo o volume

da câmara quando Ppl = Pi. Quando V aumenta, cresce também o raio do
tubo T, diminuindo a resistência hidrostática.
Para valores constantes do volume V (ou da diferença de pressão Ppl –
Pi), pode obter-se uma família de curvas do caudal F em função de Pi (Fig.
_____ II.46).
23
Fig. II.46 - Família de curvas do caudal F em função de Pi

valores constantes do volume V (ou da diferença
de pressão Ppl – Pi).
O valor máximo (caudal máximo) destas curvas aumenta com o valor

do parâmetro V, ocorrendo também aumento dos valores de Pi, para os quais
ocorrem os máximos. O primeiro destes factos resulta da diminuição de
resistência, com o aumento de V.
Sendo conhecida a Eq. II.91 que nos permite saber o valor de Ppl – Pi ,
para cada V, e usando a Fig. II.46, podemos traçar curvas de F em função de
Ppl, cujos perfis se encontram na Fig. II.47.

_____
23
Fig. II.47 - Curvas de F em função de Ppl para valores constantes do

volume V obtidas a partir das curvas da Fig. II.46.

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donde
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Para um homem de 80 Kg, D = 7,7u0,05u80 = 30,8 Kg.

A análise das forças em jogo nas articulações pode levar a conclusões
interessantes.
Os músculos esqueléticos operam entre dois ossos ligados entre si por
uma articulação. Quando o músculo (não representado na Fig.III.100) se
contrai exerce forças sobre os dois ossos5 em que se insere, forças que se
situam na mesma linha de acção, são iguais em intensidade e de sentidos
o Bo
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os ossos, ligeiramente, para evidenciar as forças que actuam na
articulação.
Por outro lado, numa situação de equilíbrio, o músculo ao exercer uma

acção de tracção sobre um osso, desencadeia forças de compressão entre as
superfícies comuns da articulação, Fig. III.100-b). Nesta figura, com fins
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o
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o o
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Notar porém que o estudo que acabámos de fazer considerou unica
mente a força muscular e a reacção na articulação. Contudo, estão também
presentes no sistema o peso dos componentes e a reacção na ligação do braço
ao ombro na articulação úmero-escapular, esta sem qualquer interferência na
articulação do cotovelo quando o braço se supõe imóvel. Pode também ocorrer
que um peso esteja a ser sustentado na mão, ou outra acção em curso.
O equilíbrio na presença de forças externas aplicadas implica
alterações nas forças compressivas a actuar na articulação.
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SÉRIE ENSINO
IMPRENSA DA UNIVERSIDADE DE COIMBRA
COIMBRA UNIVERSITY PRESS
2014

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II.2 - Lei de Boyle

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Fig. II.1 - Dispositivo para verificação da lei de Boyle.

Suponhamos que o desnível entre as superfícies livres do mercúrio em

onde Po é a pressão atmosférica, U a massa específica do mercúrio e g a ace-

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Se a pressão se mantiver constante, uma certa massa de gás aumenta

onde V é o volume a t graus centígrados e Vo o volume a 0 ºC.

Fazendo um gráfico de V em função de t obtém-se uma linha recta

onde T significa temperatura em graus Kelvin e t temperatura em graus cen-

onde K' é uma constante.

A lei de Charles pode pois enunciar-se de modo diferente:

A lei de Charles podia verificar-se utilizando o dispositivo represen-

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