.^ This is one way to allow me to continually research and develop more great products that will help you achieve your fat loss and muscle building transformation goals.
^ Tags: Building Muscle challenge coaching Energy exercise fat loss Interview lean bodies Metabolic Surge muscle building Nick Nilsson nutrition Questions supplements transformation challenge workouts .
^ If you are already active, be more active and you won’t need to adjust your calories, just your activities will increase a lean body lifestyle.
The study is therefore very largely a
study of the history of the food of the body, since it is in the
food that the necessary matter and energy are supplied.
.^ Variation in serving sizes, preparation techniques, product testing and sources of supply, as well as regional and seasonal differences may affect the nutrition values for each product.
.^ Vitamin B2 • Aids in formation of red blood cells and antibodies • Essential for carbohydrate, protein and fat metabolism • Promotes general health • Necessary for the maintenance of...- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
^ Can Calories: 0 Fat: 0g Cholesterol: 0mg Sodium: 45mg Total Carbohydrates: 0g Fiber: 0g Sugars: 0g Protein: 0g Vitamin C: 0% .- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
^ Calories: 0 Fat: 0g Cholesterol: 0mg Sodium: 25mg Total Carbohydrates: 0g Fiber: 0g Sugars: 0g Protein: 0g Vitamin C: 0% .- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
They are treated under the following heads: I. The
Chemistry of Digestion; II.
The Mode of Formation of the Digestive Secretions; III. The
Mechanism by which the Food is passed along the
Alimentary
Canal; IV. The Absorption of Food; V.
Metabolism; VI.
Excretion.
.^ I found it to be interesting because it says that it doesn't matter what kind of diet change one under goes, low carb, high carb, high protein, hig...- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
.^ As a result, it can take more than twice as long for caseins to be broken-down into their amino acid subcomponents than whey and other proteins.- Bulk Nutrition - Optimum Nutrition 100% Casein Protein at discount prices! 8 January 2010 5:19 UTC www.bulknutrition.com [Source type: Academic]
^ Glucosamine and Chondroitin are substances that are produced in our bodies naturally and help serve as building blocks for your connective tissues.- Nutrition - Find great deals, lowest prices, and buy Nutrition at Shopping.com 8 January 2010 5:19 UTC www.shopping.com [Source type: General]
^ When you suddenly cut down calories to that low an amount your body thinks it is starving and goes into surviv...- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
.^ If you don't correct under-nutrition, either your brain or some other part of your body is going to give.- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
r.
Enzyme Action
generally. - The substances which bring about this change are
known as ferments, enzymes or zymins. Formerly it was believed that
there were two distinct classes FIG. 3. - Myristica
fragrans. I. Male
flower X 2.2. Female flowerX 2.
After
Berg and Schmidt. From
Strasburger's
Lehrbuch der Botanik, by permission of
Gustav Fischer.
FIG. 2. -
Myristica fragrans, seed cut through longitudinally. (Official.) g,
Aril.
h, Outer integument, interrupted at r by the raphe.
m, Ruminated sperm.
n, Embryo (nat. size).
endo of enzymes, those which were living or associated with
living cells, and those which were non-living. In 1897, however, E.
Buchner and M. Hahn showed that from living cells (yeast) a ferment
could be obtained which acted quite as well extracellularly as when
it was bound up within the
cell.
.^ In an idealized setting, there is very little or no exchange of sediment between the pocket beach and the adjacent shorelines.- Beach Nourishment - Coastal Geology 20 November 2009 10:16 UTC www.csc.noaa.gov [Source type: Academic]
^ When doctors told me there was no more they could do, I found this site and found people who were going through the same symptoms I was going through thanks to Paxil.- All About Nutritional Supplements: Consumers Review Supplements, Vitamins, Herbs and Prescription Drugs 8 January 2010 5:19 UTC www.nutritionalsupplements.com [Source type: General]
^ In its idealized form, this type of beach is one where there is no sand present other than that resulting from beach nourishment.- Beach Nourishment - Coastal Geology 20 November 2009 10:16 UTC www.csc.noaa.gov [Source type: Academic]
All ferments probably act as
catalysators or catalysts.
Catalysis is the process by which reactions
are either initiated or accelerated by the mere presence of certain
substances which remain unchanged during the process; to these
substances the name of catalysators has been given. As an example
of such catalytic action the
acceleration of the decomposition of
hydrogen peroxide (H 2 0 2)
into water (H 2 0) and oxygen (0) by the action of a colloidal
solution of
platinum may
be given. C. Oppenheimer defines an enzyme as a substance produced
by living cells, which acts by catalysis. E. Fischer has shown that
the action of ferments is specific, that is, the ferment only
exerts its action on definite substances or substrates of definite
structural arrangement. He has compared the relation of ferment to
substrate to that of a
key to its
lock.
.^ A nutritionally sound approach to a high protein diet assists in building lean muscle without piling on fat and risking your health.- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
^ Vitamin B2 • Aids in formation of red blood cells and antibodies • Essential for carbohydrate, protein and fat metabolism • Promotes general health • Necessary for the maintenance of...- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
^ Can Calories: 0 Fat: 0g Cholesterol: 0mg Sodium: 45mg Total Carbohydrates: 0g Fiber: 0g Sugars: 0g Protein: 0g Vitamin C: 0% .- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
The chemical composition of enzymes is unknown.
.^ I found it to be interesting because it says that it doesn't matter what kind of diet change one under goes, low carb, high carb, high protein, hig...- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
In all probability the
protein is there as an impurity owing to incomplete separation.
As regards the general properties of enzymes, most of them can
be precipitated from their solutions by means of
alcohol. They can also be carried down by fine
precipitates of certain inorganic salts or by protein
precipitation,
e.g. when a precipitate of casein is
produced by acidifying a casein solution with
acetic acid. Most of
the ferments are soluble in water or saline solutions, and in
glycerin and water. The
ferments are found to have an optimum temperature of action. This
temperature in most cases ranges from 37° to 40° C. All true
ferments are thermolabile, being destroyed at about 70° C. Ferments
are hindered in their action to some extent by the general
protoplasmic poisons, such as
salicylic acid,
chloroform, &c. The action of many of
them is retarded when the products of their action are allowed to
accumulate.
.^ Thus, it is important for the stakeholder to understand that over several years the nourished beach width will decrease with the rate of this evolution decreasing and approaching a more or less equilibrium dry beach width.- Beach Nourishment - Coastal Geology 20 November 2009 10:16 UTC www.csc.noaa.gov [Source type: Academic]
^ Marion Nestle Just when the melamine scandal seemed over, new chemicals and cover-ups emerge.- The Atlantic Food Channel: Nutrition 8 January 2010 5:19 UTC food.theatlantic.com [Source type: General]
In the
case of certain enzymes at least this equilibrium may be reached
from either side; thus the enzyme maltase may either bring about
the breakdown of the
sugar
maltose to dextrose or cause a
synthesis of dextrose to maltose.
|
Material acted on.
|
Enzyme.
|
Where found.
|
|
Pepsin
|
Gastric juice
|
|
Trypsin
|
Pancreatic juice
|
|
I. Protein .
|
Erepsin
|
Small intestine
|
|
Various autolytic
enzymes
|
Tissues generally
|
|
II. Fats. .
|
I Lipase
|
Pancreatic juice and
certain tissues
|
|
Ptyalin
|
Saliva
|
|
(salivary diastase)
|
|
|
Pancreatic diastase
|
Pancreatic juice
|
|
Maltase
|
Pancreatic juice
|
|
III. Carbohydrates
|
|
Small intestine
|
|
Invertase.
|
Small intestine
|
|
Lactase
|
Small intestine
|
|
Various tissue
diastases
|
Liver, muscle, &c.
|
A number of the body ferments have now been shown to exist in
the tissues in an inactive form. This condition is known as the
proferment or zymogen state, and before any action can be exerted
it must be activated, usually by some specific substance, as in the
case of the activation of trypsinogen by means of enterokinase. The
following table gives a list of the principal ferments concerned in
the digestion and metabolism of food-stuffs: - Certain oxydases,
catalases and de-amidizing enzymes are found in the tissues
generally and play an important part in the various metabolic
processes.
2.
Digestion in the Mouth. - The first of the digestive
secretions which food comes into contact with is the saliva. This
is the mixed secretion from the various glands, salivary and other,
the ducts of which open in the mouth. The saliva, which is for the
most part produced by the three large salivary glands, the parotid,
the sub-maxillary and the sub-lingual, is a colourless or a
slightly turbid viscous fluid with a faintly alkaline reaction and
of low specific gravity. It contains a very small proportion of
solids, which vary somewhat in amount and character in the
secretions of the different glands. Mucin and traces of other
proteins are present. Small amounts of
potassium sulphocyanide may nearly always be
detected. The functions of the saliva are twofold. First, it has a
mechanical action moistening the mouth and the food and thus aiding
mastication and swallowing by securing the formation of a proper
bolus of food; it also assists by binding the particles together,
an action of special importance when the food is dry. Second, in
man and in some of the lower animals the enzyme ptyalin exerts an
action in digestion on part of the carbohydrates of the diet. The
starches or polysaccharides are broken down, first of all to the
simple dextrins and then to the still more simple disaccharide,
maltose. The further breakdown of the maltose is carried out in the
intestine by the action
of a ferment maltase which does not exist at all or only in the
merest traces in the buccal secretion. The action of ptyalin on
starches is thus very similar to that of acids, except that it
stops at the formation of maltose. Ptyalin acts best at a
temperature of about 40° C. and in a neutral or faintly alkaline
medium, its action being inhibited by the presence of even very
dilute solutions of the mineral acids. If the
acid be in sufficient amount the enzyme is
destroyed. For this reason the action ceases in the
stomach whenever the bolus is
completely permeated by the gastric juice. As it takes time for the
gastric juice thoroughly to permeate the food mass, which remains
for a considerable period in the fundus of the stomach unmixed with
the secretion, salivary digestion goes on for about half an hour
after food is taken.
3.
Gastric Digestion. - The passage of food from the
mouth to the stomach will be dealt with later.
.^ Alison E. Field Americans are replacing home-cooked meals with prepared foods--does that lead to obesity?- The Atlantic Food Channel: Nutrition 8 January 2010 5:19 UTC food.theatlantic.com [Source type: General]
But the stomach cannot be regarded as an
essential organ, since it has been removed in dogs and in man
without apparent interference with nutrition and health.
Gastric digestion is brought about by the action of the gastric
juice, a clear watery, colourless and strongly acid fluid with a
specific gravity of about 1003. The amount of solids present is
extremely small, about o 3%. They consist of protein, nucleic acid,
lecithin and inorganic salts, in addition to the more important
constituents, the enzymes and
hydrochloric acid.
The amount of hydrochloric acid present in the juice varies with
the period of digestion. In man the maximum acid concentration is
about o. 2%. The acid exists in the stomach in two forms as free
hydrochloric acid and as combined hydrochloric acid. The amount of
each depends on various factors: (1) the secretion itself; (2) the
nature of the food; and (3) the rapidity with which the stomach
empties itself, &c. For instance, after a protein-free meal the
hydrochloric acid is for the most part free, whereas, when protein
is present, it combines with it and, unless secreted in very large
amount, most of the acid is in a fixed condition.
The hydrochloric acid is formed by the activities of certain
gland cells in the middle region of the stomach, and the fact that
it does not exist as such in the blood proves that it is formed
within these cells.
.^ In its idealized form, this type of beach is one where there is no sand present other than that resulting from beach nourishment.- Beach Nourishment - Coastal Geology 20 November 2009 10:16 UTC www.csc.noaa.gov [Source type: Academic]
That the
chlorine
comes from the sodium chloride in the food has been shown by the
fact that, when the tissues are deprived of this
salt, and sodium bromide is given, hydrobromic
acid may appear in the gastric secretion.
The hydrochloric acid is essential for the action of the gastric
enzyme,
pepsin, in splitting
up the protein of the food. In addition to this, the acid has a
slight action in splitting polysaccharides and disaccharides.
Lastly, it. acts as a bactericidal
agent, preventing bacterial decomposition from
taking place, and it may thus prevent certain noxious bacteria,
taken in in the food, from gaining access to the intestinal tract,
where there is a
chance of
their flourishing in the rich alkaline medium. It is owing to the
presence of hydrochloric acid that gastric juice can be kept for
prolonged periods without undergoing putrefaction.
The quantity of juice secreted varies with the nature of the
food consumed. Thus in one experiment, after the use of a test meal
consisting of 25 grammes
bread
and 250 c.c.
tea, there was a flow
of 106 c.c., whereas in another case with an ordinary meal there
was an output of practically 600 c.c. gastric juice.
|
Hour.
|
Quantities of Juice in c.c.
|
Digestive Power in mm.
|
|
Flesh.
|
Bread.
|
Milk.
|
Flesh.
|
Bread.
|
Milk.
|
|
1st
|
11.2
|
10 6
|
4.0
|
4'94
|
6.10
|
4.21
|
|
2nd
|
II.3
|
5'4
|
8.6
|
3.03
|
7.97
|
2' 35
|
|
3rd
|
7.6
|
4.0
|
9.2
|
3.01
|
7.51
|
2.35
|
|
4th
|
5'1
|
3.4
|
7'7
|
2.87
|
6'19
|
2.65
|
|
5th
|
2.8
|
3.3
|
4.0
|
3.20
|
5.29
|
4.63
|
|
6th
|
2.2
|
2.2
|
0.5
|
3'58
|
5.72
|
6.12
|
|
7th
|
1.2
|
2
|
..
|
2.25
|
5.48
|
..
|
|
8th
|
o 6
|
2.
|
..
|
3.87
|
5.50
|
|
|
9t
|
..
|
0.9
|
|
|
5'75
|
|
|
lot
|
..
|
0.4
|
|
..
|
|
|
Pawlow has shown that not only does the amount of juice secreted
vary with the nature of the food ingested but that the digestive
activity of the secretion also varies in the same way. He gives the
following table: Quantities
and Properties of Gastric Juice
with Different Diets: 200 gms. Flesh, 200 gms. Bread, 600 c.c. Milk. Thus each separate food
gives rise to a definite hourly secretion of the juice and to a
characteristic alteration in its properties. The
meat diet brings about a very rapid flow, the
maximum output taking place within the first two hours; with bread
the maximum output is even earlier. With milk somewhat later. When
the juice is examined as regards its digestive activity, it is
found that with meat the most active juice is secreted within the
first hour, with bread in the second and third hours, and with milk
in the sixth hour.
According to the nature of the food, the stomach seems to be
stimulated to form a secretion which will best serve its purpose
and give the minimum of waste. It thus works economically.
The principal ferment found in the gastric juice is pepsin, a
ferment which acts only in the presence of a mineral acid. The
action proceeds best at a temperature of about 37° C. in an acid
medium of 0.2% to 0.3%. Pepsin is elaborated in the so-called chief
cells of the gastric glands as an inert precursor-propepsin. It is
only when it comes into contact with the acid of the juice that it
is activated and rendered capable of attacking the protein of the
food.
As already mentioned, the main function of the gastric juice is
to deal with the protein moiety of the food and to prepare it for
further digestion in the intestine.
.^ As a result, it can take more than twice as long for caseins to be broken-down into their amino acid subcomponents than whey and other proteins.- Bulk Nutrition - Optimum Nutrition 100% Casein Protein at discount prices! 8 January 2010 5:19 UTC www.bulknutrition.com [Source type: Academic]
This body may be regarded mainly as the product of the
action of the hydrochloric acid independently of the pepsin.
The following steps of decomposition are the result of the
action of pepsin. From the metaprotein primary and secondary
proteoses, the so-called proto-, heteroand deutero-albumoses are
formed, and from these peptones are finally produced.
.^ I found it to be interesting because it says that it doesn't matter what kind of diet change one under goes, low carb, high carb, high protein, hig...- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
^ When you suddenly cut down calories to that low an amount your body thinks it is starving and goes into surviv...- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
^ High Protein Diet for Weight Gain .- Diet & Nutrition | LIVESTRONG.COM 8 January 2010 5:19 UTC www.livestrong.com [Source type: General]
Formerly
it was believed that the action of the pepsin on protein could not
carry the decomposition further than the peptones, but recently it
has been shown that still further splitting can be brought about,
and that the simple amino acids of which the protein
molecule is built up can be
produced. This latter process, however, takes a very long time even
under favourable circumstances, and it probably never occurs under
normal conditions. The contents of the stomach-products of protein
digestion-are passed on into the duodenum, chiefly as proteoses and
peptones.
In addition to the principal ferment of the gastric juice some
workers hold that another enzyme is present. This is the ferment
rennet, rennin, or chymosin, the sole action of which, so far as is
known at present, is to bring about the curdling of milk, the curd
formed being dealt with in the ordinary way by the pepsin. Clotting
of milk under the action of rennin occurs at a suitable temperature
with great rapidity. This process is said to take place in two
stages: (I) the rennin converts the caseinogen of the milk into
paracasein, and (2) this paracasein unites with the
lime salts present in the milk and forms the curd
or precipitate. That lime salts are absolutely essential for this
process of clotting has been shown by the fact that, if they are
removed by precipitation as by oxalates, no clotting will take
place even after the addition of a large amount of active rennin.
Immediate clotting takes place, however, when the necessary lime
salts are restored. Many observers now hold that this rennet action
is not the property of a specific ferment but simply another phase
of the action of pepsin. For this view, which has been put forward
by wellknown workers, there is much to be said and certainly the
power of curdling milk is not confined to the stomach, but has been
found in various
tissue
extracts, and, indeed, wherever proteolytic enzymes are found.
The speed with which the stomach is emptied depends to a great
extent on the nature of the food. Plain water leaves the stomach
almost at once, salt and sugar solutions at a somewhat slower rate.
Milk under the action of rennin curdles. The whey rapidly leaves
the stomach, whereas the casein and fat are retained for further
treatment. On a mixed diet, emptying of the stomach in man proceeds
very slowly, requiring about four hours.
Cannon, by feeding with food impregnated with
bismuth and using X-rays,
showed that carbohydrates leave most rapidly, then mixtures of
carbohydrates and proteins, then proteins, then fats, and finally
mixtures of fats and proteins. The diet which remains longest in
the stomach is a mixture of fats and proteins-rich food, as it is
popularly called. Here two factors enter to prevent rapid emptying:
(I) the presence of much fat, and (2) the acid secretion engendered
by the abundant protein.
There is no doubt that fats present in fine emulsion can be
decomposed in the stomach. The action proceeds in a medium which is
slightly acid or neutral, being entirely prevented by the presence
of strong acids and alkalis. Many workers believe this gastrolipase
to be of pancreatic or intestinal origin, and suppose that it gains
entrance to the stomach by a reflux 'flow through the pylorus.
Evidence is accumulating to show that this view is correct.
By means of pepsin and gastrolipase proteins and fats are dealt
with. No specific enzyme for carbohydrates has been found in the
stomach in man. Certainly a small amount of polysaccharide
decomposition takes place, but this is dependent (I) on the ptyalin
which comes from the mouth, and (2) on a certain amount of
hydrolysis due to the action of the free hydrochloric acid.
4.
Digestion in the Intestine.-The passage of food from
the stomach to the intestine will be considered later. The food so
far digested in the stomach is known as chyme, and it is passed on
to undergo intestinal digestion under the influence of (I) the
enzymes of the
pancreas,
and (2) of other enzymes present in the different secretions of the
intestine. Digestion in the intestine may accordingly be described
under these two heads.
(
a) Pancreatic Digestion.-The pancreatic juice is the
secretion from the pancreas and is discharged into the duodenum.
The secretion obtained from a
fistula of the pancreatic duct varies in
character according to whether the opening into the duct has been
made recently or some time before the examination. It is a clear,
usually thin fluid with a specific gravity of about 1008, and with
an alkaline reaction. It contains a certain amount of protein and
ash. The most important inorganic
constituent is sodium carbonate, which gives the alkaline reaction
(alkalinity is, as NaOH = o 47%. This alkaline salt, along with
that contained in the intestinal juice, plays an important part in
neutralizing the acid chyme.
In the pancreatic secretion there are at least three important
enzymes, each with a definite action: (a) trypsin, the proteolytic
enzyme which brings about the further breakddwn of the food
proteins; (b) a diastase which deals with the
carbohydrates, and (c) a lipase which acts on the
fats.
(a) Trypsin.-This ferment, in the form in which it is
secretedtrypsinogen-is inert. Before it can exert its hydrolytic
action it must be activated. This activation is brought about by
another enzyme which is found in the intestinal tract-enterokinase.
The conversion is brought about as soon as the trypsinogen comes
into contact with the enterokinase, the merest trace of which
suffices to activate a large amount of trypsinogen.
Trypsin acts on the protein j ust as pepsin does, by bringing
about hydrolytic changes. It differs from the latter in acting best
in an alkaline or neutral medium. Its effect is much more energetic
than that of pepsin, so that the protein molecule is more
completely decomposed. Whilst it generally finishes the
decomposition which the pepsin has begun, it can break down the
original protein quite as easily if not more easily than does
pepsin, and it carries the splitting as far as the comparatively
simple crystalline bodies, the amino acids, or groups of these, the
polypeptides, bodies intermediate between the complex peptones and
the simple amino acids of which the protein is built up.
The character and properties of the products formed in such
digestion depend on the nature of the protein acted upon. As will
be seen from the following table these proteins vary fairly widely
in the proportion of amino acids which they contain.
|
Caseinogen.
|
Gelatine.
|
Globine
from Oxy-
haemoglobine.
|
Elastine.
|
|
Glycocoll
|
. .
|
16 5
|
. .
|
25.75
|
|
Alanine. .
|
0 9
|
o 8
|
4 19
|
6.58
|
|
Leucine. .
|
10.5
|
2 I
|
29.04
|
21.38
|
|
aProline. .
|
3 I
|
5.2
|
2.34
|
I'74
|
|
Phenyalanine
|
3.2
|
0.4
|
4'24
|
3'89
|
|
Glutamic acid
|
10.7
|
o 88
|
1.73
|
o 76
|
|
Aspartic acid
|
I .2
|
0.56
|
4.43
|
|
|
Cystine
|
o 06
|
. .
|
0.31
|
..
|
|
Serine. .
|
0.2
|
..
|
0.56
|
|
|
Oxyproline .
|
0.25
|
3 o
|
1.04
|
..
|
|
Tyrosine .
|
4'
|
|
I' 33
|
0.34
|
|
Lysine. .
|
5.80
|
2.75
|
4'28
|
|
|
Histidine .
|
2'59
|
0.40
|
Io 96
|
..
|
|
Arginine. .
|
4'84
|
7 62
|
5'42
|
0'3
|
|
Tryptophane
|
1-
|
..
|
Present
|
|
ioo
Grammes Protein yielded Whether any of the
polypeptides found in digestion are further broken down in the
course of normal pancreatic digestion is a
moot point, but E. Fischer and E. Abderhalden have
shown that many of the synthetic polypeptides prepared by them can
be broken into their constituents by the action of trypsin. The
previous peptic digestion seems to play some part in the extent to
which tryptic digestion is carried out, as one of these observers
has demonstrated that protein digested first with pepsin and then
with trypsin gives a smaller yield of polypeptide and a larger
yield of monamino acids than when digestion has been carried out
with trypsin alone.
b. Diastase
This ferment is found in the pancreatic juice apparently
secreted in an active form, although some observers hold that it
also is secreted in a zymogen form. It is practically identical in
its action with the ptyalin of the saliva, converting
starch into maltose. It deals
with all the starchy food which has escaped conversion into the
simple sugars by the ptyalin.
c. Lipase
Most of this ferment, if not all, is apparently secreted in the
form of a zymogen. There is evidence that the bile is the
activating agent here, just as the enterokinase acts in the case of
trypsin. Lipase can act in any medium acid, neutral, and alkaline,
and both on emulsified and non-emulsified fats. It converts the
fats by a process of hydrolysis into fatty acids and glycerin.
Kastle and Loevenhart found that not only can this enzyme break up
fats into their components, but that it also has the power to act
in the reverse direction, and in this way bring about the union of
fatty acids and glycerin so as to form fats, a process which occurs
in the intestinal epithelial cells after absorption.
In addition to these three enzymes the pancreatic juice may
contain traces of others, for example, a rennet-like ferment which
curdles milk. This again, as in the case of the stomach rennet, is
held by some to be only another phase of proteolytic action.
Maltase is also said to be present in small amount, as is also
lactase under certain conditions. In pancreatic, as in gastric
digestion, the nature of the food is said to play a part in
controlling the amount and the composition of the secretion with
respect to its ferments. The action, if it does exist, is not very
well defined.
b. Intestinal Digestion
By this is meant the other digestive processes which go on in
the intestine under the action of the secretion of Lieberkiihn's
follicles - the
succus entericus. This is a yellowish,
often opalescent, strongly alkaline fluid. The alkalinity is due to
the presence of sodium carbonate. It contains a small amount of
protein,
shed epithelial cells,
&c. The secretion of some 170 c.c. in 24 hours has been
observed in a short
loop of human
intestine by H. S. Hamburger and E. Hekma, but it is almost
impossible to get a measure of the actual amount of secretion from
the whole gut. Most of the ferments are present in very small
amount in the intestinal juice. They seem to be actually within the
epithelial lining of the intestine, for extracts made from the
intestinal mucous membrane are richer in ferments than the
secretion.
Apparently the intestinal secretion contains no trace of a
ferment acting on native protein, but a ferment - erepsin - is
present in fair amount in the intestinal mucous membrane and in
small amount in the secretion, which acts in an alkaline medium on
proteoses, peptones, and on casein, converting them into
crystalline products of the nature of amino acids.
Another ferment, arginase, has been isolated from the intestinal
mucous membrane by A. Kossel and H. D. Dakin, which splits the
diamino acid arginin into
urea and
ornithin. A lipase has also been detected which is very similar to
pancreatic lipase; it, however, attacks only emulsified fats.
Several
carbohydrate hydrolysing enzymes have been
described in the small intestine. Invertin, the ferment which
splits
cane-sugar, is present in
small amount in the secretion, more abundantly in the
extract of mucous membrane. In
all probability it deals with the saccharose after or in process of
absorption. Maltase is also present in large amount, and here again
in greater amount in the extract than in the secretion. The
presence of lactase has been much discussed, and it seems probable
that suckling animals do possess this enzyme. Some workers have
stated that an intestinal diastase is to be found, but, if so, it
is present in very small amount.
In the large intestine a small amount of erepsin has been
discovered at the upper end. Any digestion which does take place is
probably either bacterial in origin, or due to ferments which have
originated in the lower end of the small intestine, and which have
been carried down.
5. Bile
This fluid, in all probability, has little direct action in
ordinary digestion, although it contains substances which act
indirectly. The bile salts act as solvents for fats and fatty
acids, and as activators of pancreatic lipase. The salts also serve
to keep cholestrin in solution. Bile is to be looked upon rather as
the excretion, the result of the hepatic metabolism, than as a
digestive juice. Various workers have shown that when the bile is
prevented from entering the intestine owing to a fistula having
been made, the animal or patient may continue to enjoy good health,
thus proving that this fluid is not essential to any of the
digestive processes which normally take place.
Bile as secreted has an orange-brown colour, but the colour
varies according to the pigment present. It is more or less viscous
(not so viscous as bile taken from the
gall bladder) and has a specific gravity of about
Iwo. It has a slightly alkaline reaction, a bitter taste and a
characteristic
smell. The daily
output is, for a normal individual, over 500 c.c. On analysis it is
found to have over 2% of solids, of which more than half are
organic. It contains in addition to a nucleo-albumin, derived
mainly from the bile passages and gall
bladder, bile acids, bile
pigments, cholesterin, lecithin, fats, &c.
The most abundant solids are the salts of the bile acids, of which
in man the most important is sodium glycocholate, sodium
taurocholate being present in very small amount. The bile acids are
formed in the
liver cells, and
when the duct is ligatured they tend to accumulate in the
blood.
The pigments amount to only about 0.2%. In human bile the chief
pigment is bilirubin, whilst in herbivora biliverdin is more
abundant. They are derived from the haemoglobin of the blood, but
the pigments are
iron-free. They
may be regarded as purely excretory products arising from the
breakdown of the haemoglobin of effete blood corpuscles.
Cholesterin is a monatomic alcohol, and is probably a waste
product. It occurs in the bile only in small amount, and there is
some evidence that it is not secreted by the liver cells but is
added to the bile from the bile passages. Fats and lecithin are
both derived from the liver cells. Of the inorganic constituents
phosphate of
calcium is the
most abundant.
The secretion of bile is practically continuous, but it seems to
enter the duodenum intermittently. The taking of food increases the
flow of bile, the amount of the increase depending to a certain
extent on the nature of the food. A protein meal has been found to
have the greatest effect and a carbohydrate one the least. The
entry of the acid chyme into the duodenum is the stimulus which
brings about the ejection of the bile. Pressure on the liver also
seems to cause a flow (see section II.).
In connexion with bile secretion attention may be drawn here to
a peculiar enterohepatic circulation which is stated to exist. The
bile salts are partly absorbed from the intestine, to be carried
again by the portal blood to the liver and to be again eliminated.
By this circulation the entrance of various alkaloidal and ptomaine
poisons into the general circulation may be prevented.
Faeces
The bulk of the waste matter arising from the foods along with
the secretions from the alimentary canal form the faeces. On an
absorbable diet the faeces are almost purely intestinal in origin.
As a channel of excretion of nitrogenous metabolic waste products
they are not very important, although the work of C. Voit indicates
that they do play a certain part. The nature of the excreted
nitrogenous substances has not been fully examined. Of the
inorganic constituents iron is probably for the most part excreted
into the large intestine. It is, however, very difficult to come to
any definite conclusion as to what is unabsorbed material and what
excreted.
II. THE Mode Of Formation Of The Digestive Secretions 1.
Salivary Glands. - The secretion from the various glands
is generally evoked by nervous impulses, through the secretory
nerves. K. Ludwig found that the stimulation of the chorda tympani
produced a copious flow of watery saliva from the submaxillary
gland, and a general
dilatation of the blood-vessels supplying
the gland. The same is the case in the sublingual gland. In
addition to the chorda tympani
fibres also pass to the gland through the
cervical sympathetic, and when these are stimulated the saliva
excreted is viscous and turbid, and contains much solid matter,
while the blood-vessels are contracted. The conclusion formerly
drawn was that the flow of saliva was dependent on the increased
blood supply. But it has been definitely proved that true secretory
fibres exist. If atropine be administered before stimulation of the
chorda tympani, the dilatation of the vessels takes place, but no
flow of saliva. Further, if the circulation be cut off from the
gland the stimulation of the chorda tympani may cause a temporary
flow of saliva.
The parotid gland is supplied by the auriculo-temporal
nerve which receives its secreting
fibres from the glossopharyngeal. Stimulation of these fibres
brings about an abundant watery secretion poor in solids.
Stimulation of the sympathetic fibres system is not followed by any
salivary flow, yet it has an effect on the gland, for, if after the
sympathetic has been stimulated a secretion be evoked by
stimulation of the glossopharyngeal nerve, the saliva secreted is
very rich in organic solids.
2. Gastric Glands
The control of the gastric secretion seems to be under two
entirely different mechanisms. Pawlow has clearl y shown that the
stomach is supplied with secretory nerves which reach that organ
through the vagus. The stimuli which bring these nerves into action
are the sight, the odour or the taste of food. That the course of
the stimulus is through the vagus is shown by the fact that an
abundant flow of juice may be caused so long as the vagi are
intact, but this flow does not take place when these nerves are
cut. Between the stimulation and the secretion there is a lengthy
latent time amounting to several minutes. The other stimulus of the
secretion is apparently a chemical one. Pawlow states that
mechanical stimulation of the mucous membrane fails to bring about
a flow of juice, but
Beaumont in his classical observation on the
stomach of St
Martin found
that the insertion of a tube did cause a flow. There may be certain
substances either present in the food or developed in the course of
digestion, which directly stimulate the secretion originally
started by a nervous reflex. E.
Starling has drawn attention to this chemical
mode of stimulating different organs. To the substances known and
unknown which evoke the action, he gives the name of hormones, and
such "hormone" action he does not limit merely to the secretory
organs but extends to all cases where one organ is stimulated by
chemical products formed in the same or another organ. Attention
has already been drawn to the influence of different food-stuffs on
the amount and nature of the gastric secretion.
3. Pancreatic Secretion
The stimuli which evoke this secretion are again two in number.
Many have failed to demonstrate that the secretion of the pancreas
is under nervous control, but Pawlow and his school have shown that
stimulation of the vagus evokes a secretion of pancreatic juice.
This flow, as in the case of the stomach, has a latent period of
several minutes. Most modern workers hold that the most effective
stimulus to the pancreatic flow is the chemical one - a hormone
discovered by W. Bayliss and E. Starling, who found that extracts
of the duodenal mucous membrane made with dilute hydrochloric acid
when injected into the blood caused a flow of pancreatic juice. The
active substance present in this extract is known as "secretin,"
and is supposed to be formed under natural conditions by the action
of the acid chyme on a prosecretin. This secretin is not of the
ordinary zymin nature, as it is not destroyed by boiling and is
soluble in alcohol. The secretin when formed must be absorbed into
the blood and then carried round the circulation to the pancreas
before it can act.
4. Intestinal Juice
The mode of action of the stimuli which evoke this secretion has
not yet been fully investigated. As has been stated, it is quite
possible that very little ferment is secreted, and that ferment
action mainly takes place within the cells after the various
substances have been absorbed.
How far the flow is controlled by nervous action, and how far by
hormone action, is not known.
Motor Mechanism Of The Alimentary Canal
Mastication. -
This is a purely voluntary act, and consists of a great
variety of movements produced by the various muscles in connexion
with the lower
jaw. By the act of
chewing the food is thoroughly broken up and intimately mixed with
the saliva.
Deglutition
The food after thorough mastication is collected on the surface
of the tongue, principally by the action (voluntary) of the
buccinator muscles, and by the contraction of the tongue muscles it
is passed backwards. As soon as the food by the action of the
tongue enters the pillars of the
fauces the action becomes involuntary and
reflex. The soft
palate is
raised to prevent the food entering the nasal cavity, and the
larynx is shut off by
closure of the glottis, and approximation of
the
arytenoid cartilages
to one another and to the back of the epiglottis. The food is now
passed on into the
oesophagus proper by the constrictors of the
pharynx. In the oesophagus
the downward movement varies with the nature of the food swallowed.
If it be fluid it reaches the lower end of the oesophagus in about
three seconds and lies at the lower end of the gullet for two or
three seconds before entering the stomach. When the consistency is
firmer the progress downwards is much slower. Either by the force
exerted by the
wave of contraction
passing down the gullet or by some
inhibition of the sphincter, the cardiac
orifice opens and permits the food to enter the stomach.
Stomach Movements
For our knowledge of these we are indebted principally to the
work of Cannon, who studied them by feeding an animal with food
containing bismuth and then following the movements of the
shadow of the food on a
screen by means of the X-rays.
Soon after food is taken it is found that a contraction begins
somewhere about the middle of the stomach and slowly passes towards
the pylorus. This is followed by others, in man at regular
intervals of about twenty seconds, so that the pyloric part of the
organ is soon in active peristalsis. The fundus of the stomach is
not actively concerned in these movements; it simply acts as a
reservoir. At certain periods, but not with each peristaltic wave,
the pyloric sphincter relaxes and allows a portion of the fluid
acid chyme to escape into the duodenum. It only opens when
stimulated by fluid material; if solid food be forced against it it
remains tightly closed. Griitzner, by experiments with feeding with
different coloured foods, has shown that the food at the fundus may
remain undisturbed for quite prolonged periods. In this connexion
it must be remembered, of course, that the food is not lying loose
in a
sack larger than the
contents. The cavity of the stomach is only the size of the amount
of food present; in other words, the food exactly fills the cavity.
The motor nerve fibres to the stomach run in the vagi, which also
contain fibres inhibitory to the cardiac sphincter. The splanchnic
nerves mainly contain inhibitory fibres. The automatic movements
are probably in connexion with the
intrinsic plexus of Auerbach, since they
continue after section of the extrinsic nerves.
Intestinal Movements
The intestines owe their peculiar movements to the arrangement
of their muscular coats, which are disposed in two layers, an inner
circular, and an outer longitudinal. The movements are of two
kinds, the so-called swaying myogenic contraction and the
peristaltic waves. The former are rapid and have very little to do
with the downward movement of the contents. Probably their action
is to mix the contents, since Cannon has shown that these contents,
in the lower animals at least, get divided into segments. From time
to time the separated segments are caught in the course of a
peristaltic wave and carried downward a short distance. Then again
in their new situation the rhythmic contractions break up the
contents anew.
The peristaltic movements are much more powerful. Under normal
conditions they begin at the pylorus and passing downwards carry
the intestinal contents onwards. The normal movement progresses
slowly, although under abnormal conditions peristaltic waves may
become extremely violent and rapid, and may indeed run over the
whole length of the intestine within a minute. The muscular coat in
front of the contracting zone is relaxed, as is that behind the
wave. The waves are probably due mainly to the circular fibres, the
longitudinal pulling the gut up over the contents as they are
forced onwards. The downward movement seems to be due to some
definite arrangement within the intestinal wall, since it has been
shown that, when a segment of bowel has been cut out and then the
continuity of the canal made good by fixing the section so that the
lower end of the excised portion is fixed to the upper divided end
of the real gut, upward peristalsis takes place in this segment. An
anti-peristalsis has been described in which the movements are all
towards the stomach. Under certain conditions the introduction of
foreign substances, as hairs, &c., may evoke such
anti-peristaltic waves.
The rhythmical movements are held by some to be purely myogenic
in origin, as they still continue after section of all the nerves
and when the intrinsic ganglia in the intestinal wall have been
thrown out of action by the application of
nicotine. But recent work by R. Magnus would
tend to show that they are controlled by Auerbach's plexus.
Peristaltic waves, on the other hand, according to W. Bayliss and
E. Starling, although they continue and indeed may become more
energetic after section of the extrinsic nerves, are prevented by
the application of nicotine and
cocaine; in other words, it is presumed that
peristalsis is a complicated reflex action through the intrinsic
ganglia. The intestines are therefore not dependent for their
movement on their connexion with the central
nervous system,
although of course their activity is more or less regulated by such
a connexion.
As regards the movements of the large intestine, they resemble
those of the small, although they are much less frequent. The
forward movement is slow, thus permitting of the solidification of
the contents by the removal of the water. In the first part of the
large intestine anti-peristaltic movements are frequent, the
regular peristaltic downward movements only becoming prominent when
the descending
colon is reached
to carry contents to the rectum. The anti-peristalsis serves a
useful purpose in giving time for the absorption of the fluid in
the formation of faeces. The rate at which the contents travel
along the intestine varies greatly. Under average conditions the
food
residue reaches the
ileo-caecal
valve between the
small and large intestine at about four to four and a half hours
after a meal, while it takes nine hours to reach the splenic
flexure of the colon.
Defaecation
Food residues, cellular debris and substances derived from the
various secretions of the gastro-intestinal tract are forced
downwards by peristalsis, and eventually reach the rectum and
accumulate there as the faeces. The pressure of the solid and
semisolid mass gives rise to a definite sensation and a desire to
empty the rectum. The faeces are retained within the canal partly
by the horizontal direction of the rectum before it opens into the
anal canal, and partly by the action of two sphincter muscles. At
the act of defaecation the strong internal sphincter is first of
all relaxed, but unless the rectal stimulus is very strong, the
external can be kept contracted, as it is to a certain extent,
under the control of the will. The act of defaecation normally is
partly voluntary and partly involuntary. The voluntary part
consists in the contraction of the abdominal muscles, the closure
of the glottis, and the relaxation of the external sphincter and of
the levator
ani muscle, thus
allowing the horizontal part of the rectum to become more vertical;
the involuntary in the energetic contractions of the muscular walls
of the colon and rectum which sweep the contents of the whole colon
downwards. There is a centre in the lumbar enlargement of the
spinal cord which
presides over the sphincter muscles and probably over the whole
involuntary mechanism of defaecation.
Vomiting
Sometimes the gastric contents are ejected through the cardiac
opening of the stomach instead of through the pylorus. The act is a
reflex one, probably originally protective in nature, irritation of
the gastric mucous membrane being the most frequent cause. The act
is generally preceded by a feeling of
nausea and a copious salivation, succeeded by a
series of powerful expiratory efforts with the glottis closed. The
diaphragm is held firmly
contracted, then a convulsive contraction of the abdominal muscles
with a simultaneous opening of the cardiac orifice of the stomach
brings about the sudden ejection of the contents. The wall of the
stomach may also contract and press upon the contents. During the
act the glottis is firmly closed, and at the same time, if the act
be not too 925 violent, the gastric contents are prevented from
entering the nasal cavity by the contraction of the soft
palate.
IV. Absorption Mouth. - No absorption of food-stuffs
takes place here Stomach. - Absorption from the stomach occurs only
to a small extent. Water passes rapidly through the stomach and is
practically unabsorbed. Salts are apparently absorbed in a limited
amount from their watery solution, the extent of absorption
depending to some extent on the concentration of the solution.
Sugar is also absorbed to a small extent from its solutions, the
greater the concentration the greater being the amount of sugar
taken up. Alcohol is readily absorbed from the stomach. A small
amount of the products of protein digestion may be absorbed. There
is no evidence that fats are absorbed under any conditions in the
stomach.
Intestine
The greatest absorption of the foods takes place in the
intestine, especially in the small intestine. It has been shown
that over 85% of the protein has disappeared before the lower end
of the small intestine is reached. How does the absorption take
place? There are two channels for the removal of the material from
the intestine: (r) the blood capillaries spread in the villi, and
(2) the lacteals also present in the viii. The foods may reach the
blood direct or through the various
lymph channels into the thoracic duct and finally
into the blood. The lacteals of the villi are channels for the
absorption of the fatty parts of the food. The products of the
digestion of the proteins and carbohydrates reach the body directly
through the capillaries via the portal system.
Can absorption be explained by the ordinary laws of
diffusion and osmosis, or
are there certain selective activities of the living epithelial
lining ? The work of R. Heidenhain, E.
Weymouth Reid, and others shows clearly that
whatever part the physical laws play in this exchange, there are
other activities also at work. For instance, an animal's own serum
can be readily absorbed from its intestine, as can also salt and
other solutions of higher concentration than that of the blood.
Such absorption cannot be explained by ordinary physical laws. In
all such cases of absorption the epithelial lining of the gut must
be intact and uninjured. 0. Cohnheim and others have shown that
when the epithelial lining is damaged or destroyed, the intestinal
wall behaves like any other animal membrane, and the physical laws.
governing osmotic pressure come into play. Whether the nervous.
system plays any part in this absorption is not yet determined.
The form in which the various products resulting from digestion
are absorbed must next be considered.
Carbohydrates
These reach the body, as already mentioned, by way of the blood,
and in the form of monosaccharides or simple sugars. F. Rohmann
found that the absorption of the disaccharides is dependent on the
invert ferment action, and not upon their osmotic characters. E.
Weinland too has shown that if lactose be put into a lactase-free
intestine, no absorption takes place, the lactose gradually
disappearing under bacterial action, whereas when the ferment
lactase is present
glucose
and galactose the products of its splitting are absorbed as readily
as cane-sugar and maltose. E. Voit has also demonstrated the fact
that the body deals with its carbohydrate supply in the form of
mono-saccharides. He injected solutions of various sugars, monoand
di-saccharides, and found that the simple sugars were retained,
whereas the double sugars were excreted in the urine. The only
di-saccharide which can be dealt with in the body is maltose, as
there is a maltase present in the blood which splits it.
Carbohydrates which are not absorbed from the intestine are
disposed of by bacterial action, giving rise to various fatty
acids,
carbon dioxide,
&c.
Fats. -Fats are absorbed from the intestine in the form
of fatty acids and glycerin; i.e. in the form in which
they exist after the action of the lipase. That a resynthesis takes
place in the epithelium is shown by the fact that fatty acids are
of equal value with fat as a source of energy, and that as fat
absorption goes on fat droplets are seen to grow in the protoplasm
away from the free margin of the cells. As already mentioned, the
fat is removed by the lacteals from the cells to the thoracic duct,
and then to the general circulation. A small amount of the fat may
pass into the body via the blood, but this is practically all
retained by the liver. The amount of fat absorbed depends a good
deal on the nature of the fat, especially with reference to its
melting-point, fats of low melting-point being most readily taken
up.
Protein
The older workers held that the protein was absorbed in the form
of proteose and peptone. In support of this it was stated that both
proteoses and peptones could be detected in the blood stream. The
result of the most recent work tends to show that the material is
absorbed in the form of the amino acids either simple or in complex
groups, the polypeptides, and that if proteoses or peptones be
absorbed they are attacked by the
intra-cellular enzyme erepsin, which breaks them
down into the simpler products as soon as they are within the
intestinal mucous membrane. Certain proteins appear to be absorbed
unchanged; for instance, blood serum disappears from the intestine
without apparently any change through zymin attack. This fact is
made use of in practical
medicine, as, when administration of food by
the mouth is impossible, patients are frequently kept alive by the
giving of nutrient enemata. That the food thus given is absorbed is
shown by the increase of
nitrogen excretion in the urine.
In the large intestine very little absorption of nutrient matter
takes place under normal conditions, mainly of course because most
of the absorbable material is removed whilst the food is in the
small intestine. That protein matter can be absorbed is shown by
the above statement regarding nutrient enemata. The principal
substance absorbed here is water; and thus the excreta become firm
and formed.
V.
Metabolism In
all living matter there is a constant
cycle of chemical changes going on, a constant
breaking down (catabolism), and a correspondingly constant building
up (anabolism). Unless the former is covered by the latter wasting
and finally death must supervene. These two changes together make
up the metabolism, and the study of this involves a study
of the fate of the food absorbed both when it is used immediately
and after it has been stored in the tissues of the body. Protein
matter is undoubtedly the main constituent of protoplasm, but in
what form it exists there is absolutely unknown. One thing is
certain, that for the maintenance
of life a constant
supply of protein matter is necessary. In fact it might be
said that this
is the essential food and keeps the body
alive, fats and carbohydrates being merely subsidiary. In the
mammalian organism with which we are specially concerned a supply
of these latter substances
is also necessary to yield the
energy required. The amounts of these various food stuffs which
should be present in a suitable diet are dealt with under
Dietetics. Here we are only
concerned with the part played by the different materials in the
various chemical changes which are the basis of vital activity.
Not many years ago physiologists were very much in the position
of unskilled labourers who saw loads of heterogeneous material
being "dumped" for building purposes, but who did not know for what
particular purpose each individual substance was used. Thanks,
however, to the brilliant work of E. Fischer we are no longer in
this position. Gradually our knowledge is being broadened by actual
facts obtained by direct experiment, or by inference from previous
experiments. But it is still far from complete. It
is only possible to outline what is at present
known about the part played by the different food constituents in
metabolism.
Proteins
Since these alone contain the nitrogen necessary for the
building up and repair of the tissues they are essential and will
be dealt with first. In considering the digestion of proteins it
was shown that in all probability all protein food was reduced in
the intestine to comparatively simple crystalline bodies.
.^ Diet & Weight Loss .
In
addition to these acids abundant carbohydrates and fats were given.
It has since been shown that the presence of carbohydrate a certain
amount of is absolutely essential before utilization of the amino
acids can take place. Further, it has been demonstrated that only a
mere fraction of the total amino acids resulting from pancreatic
digestion is sufficient as the source of nitrogen supply for the
animal organism. Not only so, but, in spite of the attempt to
insist on the polypeptides as being the valuable nuclei for the
rebuilding up of protein in the body, it has been shown that
mixtures of amino acids from which the polypeptides have been
removed can serve as the nitrogen supply.
What then does the body gain by breaking down food material to
such simple bodies, if it is immediately to be resynthesized ?
This complete breakdown appears to be to facilitate rebuilding. The
protein in the protoplasm of each animal is characteristic and to
build up these different proteins the material must be separated
into its nuclei. An experiment carried out by E. Abderhalden shows
this very clearly. A protein gliadin absolutely different in
constitution from the proteins of blood plasma was fed to an animal
from which much of its blood had been removed, so that an active
reformation had to take place. The question to be solved was
whether by feeding with a protein so absolutely different in
constitution the nature of the freshly forming serum protein could
be radically changed. But the newly-formed serum was found to be
exactly the same in constitution as the old. The tissues had
selected simply those nuclei of the gliadin which were required and
had rejected the others.
In addition to this breakdown of protein in the intestine,
another factor of importance comes into play. After absorption from
the lumen of the gut the amino acids are not wholly conveyed as
such by the portal blood to the liver. That the portal blood
contains a greater amount of
ammonia than the systemic blood has long been
known, and Jacoby and Lang have shown that many tissues, and among
them the intestinal tissues, are able to split off from the amino
acids their amino group NH 2. Thus it would seem probable that any
excess of the amino acids formed does not reach the liver as such
but denitrified as members of the fatty acid series. The ammonia
split off is also conveyed to the liver and is excreted for the
most part as urea, within the first few hours after a protein meal.
Thus, in all probability very early after absorption and before the
products of digestion enter into combination or any synthesis
occurs, all excess of the absorbed nitrogen is disposed of. The
rest of the products circulate in the blood, yielding to the cells
the materials of which they are in need. On the other hand some
investigators still hold that resynthesis into a neutral protein
like serum albumin takes place in the intestinal wall immediately
after absorption of the
digest
products. That the leucocytes play an important part in carrying
the products of protein digestion to the tissues is indicated by
the enormous increase in their number which occurs during the
digestion and absorption of protein foods. How they act, whether
simply as carriers of the products of protein digestion combined or
uncombined, and how they give the material to the tissues is
unknown.
Carbohydrates are generally assumed simply to serve the
purpose of yielding energy in their
combustion to CO 2 and H 2 O, and to act as
protein sparers,
i.e. they save the ingestion of large
amounts of costly protein material as a source of energy. There
may, however, be other activities in which the ingested sugars play
a part, for instance, in the utilization of the nitrogen of
proteins. It has already been indicated that the nitrogen in the
products of pancreatic digestion can be used only when a sufficient
amount of carbohydrates is given at the same time. Only
carbohydrates seem to be able to do this, for it has been found
that when isodynamic amounts of fat are given the utilization does
not take place.
When taken into the body in excess of the immediate requirements
the sugar is not utilized all at once, but any excess is stored in
the form of glycogen both in the liver and the muscles. This
glycogen is an insoluble polysaccharide, and is only utilized
according to the requirements of the body, especially during
muscular exertion. Carbohydrates, when taken in in excess, are also
stored in the tissues in the form of fat. This was demonstrated by
the feeding experiments of Lawes and Gilbert at Rothamstead. They
took two young pigs of a
litter, killed and analysed one, then fed the
other for a definite time upon food of known composition,
determining the amount of protein absorbed by analysing the urine
and the faeces. They then killed the
pig and by analysis ascertained the amount of fat
put on. They found that this was far in excess of the amount of the
protein of the food which had been absorbed and was also in excess
of what could have been formed from the small amount of fat in the
food. The fat must therefore have been formed from the
carbohydrates of the food. The
consumption of larger amounts of sugar than
can be used or stored as glycogen results in its passing straight
through the body and being excreted in the urine. This condition is
known as alimentary glycosuria. The power of using and storing
sugar varies greatly in different individuals and in the same
individual at different times.
Fats
The fats simply serve as stores of energy. After ingestion, if
in small amount, they are, like carbohydrates, oxidized to the same
final products C02, and H 2 O. If in larger amount they are stored
as fat, to serve as a reserve in case of need, in the body tissues.
Like the carbohydrates they serve as the sources of part of the
energy dissipated as heat, but they are not so efficient as sparers
of protein material, evidently in part at least because they are
less easily digested and absorbed.
Factors which influence Normal Metabolism. Fasting. - During fasting
the body draws upon its own reserve of stored material for the
requirements in the production of energy, and the rate of breakdown
varies with the energy requirements. An individual who is kept warm
in
bed therefore stands fasting
longer than one who is compelled to take exercise in a cold place.
The breakdown of tissue during the early days of a fast is much
greater than later, for as the fast progresses the body becomes
more economical in its utilization of tissue. During a fast the
tissues do not all waste at an equal rate; those which are not
essential are utilized at a much greater rate than those which are
essential to the maintenance of the organism. For instance, it has
been shown that during a fast the skeletal muscles may lose over
40% of their weight, whereas an essential organ like the
heart loses only some 3%.
The essential tissues obtain their nourishment from the less
essential probably by ferment 'action, a process which has been
termed autolysis. The autolytic products of the stored material in
the tissues are practically identical with those which arise during
the ordinary gastro-intestinal digestion.
2. Muscular Work
The muscular tissue plays the most important part in general
metabolism. Not only is muscle the most abundant tissue present,
but it is constantly active and is the great energyliberating
machine of the body. Formerly it was believed on the authority of
Liebig that muscular work was done at the expense of the protein
material, but it has been conclusively shown that the real source
of energy in moderate work is the non-protein material,
carbohydrates and fats; of these the former plays the greater part
in a man on ordinary diet. If, however, the supply of
non-nitrogenous material be insufficient, then the energy has to be
supplied by the protein and the output of nitrogen is thus
increased. Variations in the amount of creatinin and
uric acid (both products of
muscle metabolism) excreted have been described. In hard work it is
sometimes found that there may be no immediate rise in the nitrogen
output on the day of the work, but that an increase is
manifest on the second or
third day after. While the excretion of nitrogen shows no increase
proportionate to the work done, the output of carbon dioxide
produced by the combustion of the carbohydrates and of the fats is
increased proportionately to the work done.
3. Internal Secretions
Evidence is accumulating to show that the activities of the
various tissues of the body are presided over and controlled not
merely by the action of the nervous system but also by chemical
substances, the result of the activity of certain organs. To these
chemical substances, as already stated, the name of hormones has
been given.
The hormone which has been most thoroughly investigated is
adrenalin, a perfectly definite chemical compound
consisting of a secondary alcohol linked to a
benzene ring. It is a product of the central or
medullary part of the suprarenal bodies. The medullary part of
these organs is developed from the sympathetic part of the nervous
system, and adrenalin acts as a stimulant to the terminations of
the sympathetic nerves which spring from the thoracoabdominal
region. These nerves control the small
arteries, and the main action of adrenalin is
to cause a powerful contraction of these vessels, and as a result a
great rise in the arterial blood pressure. For this purpose it is
now largely used in medicine. The constant supply of adrenalin in
small quantities seems to play an important part in keeping up the
tone of the blood vessels, and when, as a result of disease of the
suprarenals, the supply is cut off a serious
train of symptoms supervenes.
Allied to adrenalin is a hormone derived from the pituitary
body. This also causes a constriction of the small arteries
except those of the kidney, which it dilates. An increased flow of
urine is produced.
In the
thyroid
gland a substance,
iodothyrin, is constantly being
produced, and this appears to exercise a stimulating action on the
rate of chemical exchange in the various tissues. Under its
administration the waste of both proteins and fats is increased.
When the thyroid is removed or destroyed by disease a condition of
decreased chemical change and mental sluggishness results,
accompanied often by nervous tremors.
A difficulty in explaining these symptoms is caused by the fact
that in the thyroid are imbedded four small parathyroids, and it is
possible that these produce a special hormone. It has been
suggested that this exercises a particular influence upon the
nervous system, but further evidence is wanting.
The well-known effects of removal of the ovaries or
testes on the development and character of an animal is
due to the absence of the special hormone or hormones of these
structures. These hormones appear to be produced, in the case of
the testes at least, not in the true genital cells, but in the
intermediate cells, since it has been found that ligature of the
duct, which leads to destruction of the genital cells, does not
abolish the development of the sexual characters of the animal.
There is growing evidence that from the ovaries different
hormones may be produced in varying amounts which play an important
part in regulating the phenomena of sexual life.
The
thymus gland is a structure lying in the front of
the neck, which is best developed at the time of birth, grows very
slowly after birth, and atrophies when the age of
puberty is reached. In
castrated male animals it continues to grow and persists throughout
life. There is some evidence that it may exercise some effect upon
the growth of the testes, probably by hormone action.
Pancreas
Within recent years it has been shown that the internal
secretion of this organ plays a very important part in the
metabolism of sugar. When the organ is completely extirpated the
animal becomes diabetic, i.e. sugar appears in the urine
and the animal emaciates. How the internal secretion effects the
combustion of the sugar is not yet known. Some workers hold that
the action of the pancreatic internal secretion is to control the
sugar formation in the various sugar-forming organs, of which the
liver is the chief, others that it dominates the utilization of
sugar as a source of energy by the muscles.
These are some of the best-known examples of the way in which
the products of the activity of one organ modify the functions of
other organs. In all probability many more examples of hormone
action will be discovered, and it will be found that it plays
probably even a more important part than the nervous system in the
coordination of function in the animal.
Other factors, besides these already dealt with, play a part in
modifying the various metabolic processes, as age, temperature,
climate, &c. Very little, however, is definitely known about
these various factors.
Water and inorganic salts are quite as essential for the
well-being of the body as the energy-yielding proteins,
carbohydrates and fats. They, however, probably undergo little or
no change in the body; they are excreted pretty much in the same
form in which they are ingested. Although they are not subjected to
any very great change yet they are of immense importance. No animal
tissue can carry on its work in the absence of the various salts.
Many experiments have been carried out in which animals have been
fed on food as free from salts as possible, and, although the food
was much in excess of the energy requirements, yet all these
animals died, whereas other animals to which similar food with
salts was given throve well. The most important acids are
hydrochloric and phosphoric, and the most important bases sodium of
potassium. Calcium and
magnesium are also of importance, especially
where
bone formation is taking
place. Another element of really vital importance is iron, which is
required for the formation of haemoglobin.
VI.
Excretion While
we know comparatively little of the intermediate stages in the
breakdown of the food constituents, and more particularly of the
protein moiety, our knowledge of the final products of the
metabolic changes excreted is fairly full. The urine is the main
channel of excretion for the nitrogenous waste products. C02,
arising for the most part frdm the metabolism of carbohydrates and
fats, is excreted mainly through the lungs. Water is excreted by
the lungs, the kidneys and the skin.
So far no entirely satisfactory explanation has been given of
how a fluid like urine, having an acid reaction and containing
about one hundred times as much urea and generally more than twice
as much sodium chloride as the blood, is formed in the kidneys. The
urine is a yellowish fluid which varies greatly in its depth of
colour, from pale
amber to a
deep brown. It has a specific gravity of about 1020, varying with
the percentage of solids in solution, and it usually has an acid
reaction. It is a fluid of complex character, containing, as
already mentioned, practically all the waste nitrogen of the body.
Among the principal organic substances present are urea, ammonia,
purins (uric acid and the so-called
purin bases, xanthin, &c.), creatinin,
conjugated sulphates, various aromatic bodies and many other
substances in small amount, together with the water and inorganic
salts.
The following table from Folin gives a good idea of the average
composition of the urine as regards the nitrogen-containing
constituents, and its variation according to the nature of the diet
when this is free of creatin creatinin and the precursors of the
purins: - Urea, which forms the chief nitrogenous
constituent, amounting on an ordinary diet to about 30 grms. per
diem, is for the most part formed in the liver, from ammonia
obtained either directly from the blood after absorption from the
intestine, or resulting from the denitrification of the amino
acids. It may also arise in part from the diamino acids and from
uric acid.
Ammonia is present in the form of ammonium salts, and
forms about 4% of the total urinary nitrogen. It may exceed this
amount under certain conditions, for the most part pathological.
The ammonia is utilized by the body to neutralize acids which arise
during the various metabolic processes.
|
Nitrogen-rich Diet.
|
Nitrogen-poor Diet.
|
|
Total nitrogen.. .
|
14.8-18.2 grms. per day
|
4.8- 8 o grms. per day
|
|
Urea nitrogen.. .. .
|
86.3-89.4% of total
|
62.0-80.4% of total
|
|
Ammonia nitrogen.. .
|
3.3- 5.1%
|
4.2-11.7% „
|
|
Creatinin nitrogen
|
3.2- 4.5% ,,
|
5.5-11-1% „
|
|
Uric acid nitrogen.. .
|
0.5- I o % '„
|
I.2- 2.4% „
|
|
Undetermined nitrogen. .
|
2.7- 5.3%
|
4.8-14.6% „
|
Purins (uric acid, xanthin, hypoxanthin, &c.) are
all members of a series which have as their common
nucleus a body which E. Fischer
called purin. The most important member of this series is uric
acid. It forms about 2% of the total urinary nitrogen. Recent work
has shown that it has two quite definite sources of origin: (I)
from ingested food containing the precursors, and (2) from the
tissue metabolism. The first is known as the exogenous source, anct
the second as the endogenous. This acid is chemically known as
trioxy-purin, and may be regarded as the union of two urea
molecules with a three-carbon chain fatty acid. All the uric acid
formed in the body is not excreted as such, part being, as already
mentioned, converted into urea. The amount which is converted into
urea varies with the species of animal. In man, Burian and Schur
state that one half of the total amount is so converted. Some
workers, like Wiener, hold that uric acid may be synthesized in the
body, but while this is undoubtedly so in the case of the
bird, in the mammal it has not been
definitely established. The other chief purin bodies present in
urine are xanthin and hypoxanthin, purins less oxidized than uric
acid; the first is a dioxypurin, and the second is a monoxypurin.
The main source of total purin supply would seem to be muscle
metabolism. The mother substances from which all are derived in the
body are the nucleins. These complex bodies are apparently first
broken down by enzyme action to aminopurins. These in their turn
have their amino groups split off, and then, according to the
degree of oxidation, the different purin bodies are formed.
Creatinin
The physiological significance of this substance is as yet
unknown. The daily excretion varies little with the character of
the diet, provided, of course, that the diet be creatin creatinin
free. It appears to be proportional to the muscular development and
muscular activity of the individual. Hence it would seem to be
derived from the creatin of muscle, a substance which is very
readily changed into creatinin outside the body. In the body the
conversion of creatin into creatinin seems to be strictly limited,
and hence when creatin is taken in flesh in the food it tends to
appear as such in the urine. It would seem that it is either in
great part decomposed in the body into what we do not at present
know or that, as suggested by Folin, it may be used as a
specialized food. Whatever its source, after urea and ammonia it is
one of the most important nitrogenous substances excreted, the
daily excretion being about 1.5 grms.
The
sulphur
excreted in the urine comes chiefly from the sulphur of the protein
molecule. It is excreted in various forms. (1) As the ordinary
preformed sulphates, that is, sulphur in the form of
sulphuric acid
combined with the ordinary bases. (2) As ethereal sulphates, that
is, in combination with various aromatic substances like phenol,
indol, &c. (3) In the form of so-called neutral sulphur in such
substances as cystin, which are intermediate products in the
complete oxidation of sulphur.
Phosphorus
appears linked to the alkalis and
alkaline earths as phosphoric acid. A
very small part of the phosphoric acid may be eliminated in organic
combination such as the glycero-
phosphates, &c.
Sodium (mostly as sodium chloride), potassium, calcium and
magnesium are the common bases present in the urine.
The lungs are the important channel of excretion for
the waste product of carbon metabolism CO 2 (see RESPIRATORY
SYSTEM); and also a very important channel for the excretion of
water. As regards the skin, the sweat carries off a large amount of
the water, but it is difficult to determine the total amount. It
has been estimated that about 500 c.c. is excreted per diem under
normal conditions. Sweat contains salts, chiefly sodium chloride,
and organic waste products. Of the organic solids excreted from
this source urea forms the most important under normal conditions.
Under pathological conditions, especially when there is
interference with free renal action, the amount of nitrogenous
waste excreted may become quite important. There is also a small
amount of CO 2 excreted by this channel. (D. N. P.; E. P. C.)