MINERAL NUTRITION OF PLANTS*

By F. G. GREGORY
Research Institute of Plant Physiology
Imperial College of Science and TechHology, London, England
The range of subjects covered by the title is so great that all
aspects cannot be dealt with in the space allotted. Reference may be
made to several reviews: the absorption of electrolytes by Osterhout
(33) , the efects of rarer elements by Brenchley (7), and a review by
Pirschle (36) which covers the efects of essential elements (potas
sium, calcium, magnesium, sodium), some of the accessory elements,
and the relation of iron to chlorosis. A summary of the literature of
ammonium and nitrate nutrition is given by Pardo (35). Attention
in this review will be confned mainly to the physiological efects of
Plineral defciency. "ork on this problem has been proceeding for
some years in the laboratory of the reviewer, and as many of the re
sults are still unpublished, salient features of the research are here
presented. In so far as recent published work of other investigators
deals with this aspect of the problem it is here discussed, though the
list may not be complete.
Reference to the work of the author and his collaborators has
already been made by Steward (51), and the importance of the
method of approach is there stressed .. It is clearly desirable to main
tain cultural conditions constant so far as external factors are con
cerned. This is the method employed by Gassner & Goeze (15, 16)
and is highly commended by Steward. This method, namely, of
maintaining all factors at constant level with the exception of the
one studied, is capable of yielding precise information, but is neces
sarily limited to the actual conditions of the experiment. As the aim
of all investigations is to make generalisations of wide application,
the consequences of change in one factor should be studied at many
levels of the other factors; in a word, the interaction of factors is of
equal importance to the study of single factors in isolation. It is of
course necessary to employ statistical methods for evaluation of the
signifcance of the single factors and their interactions, but the time
is past for putting forward results without statistical estimates of
signifcance. The results of Gassner & Goeze establish clearly enough
the importance of interactions, but the adverse criticism by these
* Received February 13, 1937.
557
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Further
ANNUAL
REVIEWS
558 ANNUAL REVIEW OF BIOCHEMISTRY
authors of the work of the reviewer and his collaborators on the
grounds that interactions were not studied was premature.
Method of study.-A pure line barley (Plumage Archer) has been
used throughout the work, grown in sand culture in the open. In the
preliminary work (21) one defciency level of nitrogen, phosphorus,
and potassium was studied, but the work was immediately extended to
cover a wider range. The standard manuring used maintains excel
lent growth and gives high yields. The relative proportions of nitro
gen, phosphorus, and potassium are not arbitrarily fxed but corre
spond roughly with the proportions of these nutrients in the grain
of barley. The proportions of N: K20: P205 are maintained at 3: 2: 1.
Five levels of defciency for each nutrient have been used, each mem
ber of a series receiving one-third that of the preceding. Thus the
range of concentrations for nitrogen are as follows (per pot of three
plants): Nl, 50 mg ; N2, 167 mg ; Na, 56 mg; N, 19 mg; Ns, 6 mg.
For K20 and P205 two-thirds and one-third, respectively, are given.
The sufxes therefore correspond with increasing defciencies of
nutrient.
In interaction experiments twenty-fve combinations have been
used. Thus completC interaction experiment with nitrogen and po
tassium may be represented as follows:
N1K1(1:1) N1K2(1:lj3) N1KaCl:1/9) N1K4(1:1/27) N1K5(1;1/81)
N
2K10/3:1) N2
K
2Cl
/
3:1
/
3) N2Kg N2K4 N2K5
N
a
K1(l
/9
: 1)
NaK
2
NaKg(l/9: 1/9) NaK4 N
s
K

N4
K1(1/27: 1) N,K2 N4Ka N4K-(1/27: 1/27) N4K
G
N3Kl (1/81: 1) N5 N
e
K
a
N
6K4
N6Ks(1/81: liB
Each edge of a square, defned by N lK1, N lKo N oK., and N oK1
as the corners, would represent a series of increasing concentration of
one nutrirnt (limiting series), all other nutrients being present at
constant level. A diagonal would represent a s
imultaneous increase
in both nutrients (balanced series). The numerals in parentheses
(shown above) represent the relati.ve proportions of the nutrients in
terms of the standard amounts given.
General considerations.-To assess the efects of nutrients on
growth, eforts should be made to Cxpress such relations in numerical
terms. The chracteristics of growth chosen should throw light on
some fundamental physiological process and furnish data capable of
further analysis. The characters chosen are + (a) the rate of produc
tion and total numbCr of tillers throughout the life cycle, which is
direct measure of meristematic activity ; (b) the rate of production
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MINERAL NUTRITION OF PLANTS
559
of leaves, which is a measure of diferentiation; (c) the growth of the
leaf surface, which by further analysis allows the net assimilation rate
to be estimated; and (d) the rate of increase and fnal dry weight of
the whole plant and its organs. The plants are sampled at various inter
vals during growth and the material obtained is anal
y
sed chemicall
y
to ascertain the amounts and rates of uptake of the nutrients. By this
means numerical records of development are obtained. Further analy
sis of such data is dealt with by Gregory (17). A study of the inter
action of water and nitrogen on these lines has been published by
Crowther (10). Recently two papers on physiological ontogeny have
appeared by Ballard & Petrie (5) , and Williams (57) , which deal
with nitrogen nutrition of wheat and phosphorus nutrition of oats.
A full study with barley of the efects on growth of nitrogen and po
tassium and their interactions has been made by Mathur (29) and of
the interaction of phosphorus and potassium by Verma (53) using
twenty-fve combinations of the nutrients as described above. Some
typical results are given below (see also Tables I to IV) .
TABLE I
MAXIMUM LEAF AREA PER PLANT IN SQ. eM., 1931 [MATHUR (2)]
K
,
Nl ...... ........ 2358
N2 ...... .... .... 724
N. . ...... ... .... 252
N4 ..... . .. . ... . . 107
N5 .... .......... 69
K.
1518
699
TABLE II
K. K. K.
1056 736 716
183
84
59
MAXIMUM TILLER NUMBER PER PLANT, 1931 [MATHUR (29)]

Nl .............. 12.8 12.9 13.2 14.6 12.5
N2 .............. 8.9 8.4
N. ... . . . . . ...... 4.8 5.0
N4 ... .......... . 3.1 2.6
N5 ... ........... 1.7 1.7
TABLE III
MAXIMUM LEAF AREA
'
PER
P
LANT IN SQ. eM., 1933 [VERMA (53)]
K
l K
K, K,
K
P
l
. ..... ..... ... 1524 925
P
2 .... ..... .... . 910 685 713 528
p
..... . ..... ... 507 401 32 315 331
P4 .. . .. ... ... . .. 307 180 194 130
P5 . . . . .. . ... ... . 258 72 65
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56 ANNUAL REVIEW OF BIOCHEMISTRY
TABLE IV
MAXl.MUM TILLER NUMBER X
P
LAN'f, 1933 [VERMA (53)]
K,
]
K. K. K.
PI
.... ... ..... . . ,!o.o 21.2 24.7 24.0 14.2
P
2
.. . ...... ..... :4.4 14.8 17.4 13.2 9.5
Pa
.. ... ......... 10.8 9.3 8.8 8.3 7.8
P4 .. . .. .. . , . .... . 5.8 5,,1 4.8 3.9 3.6
P
6
.. .. . ... " ... .. 2.6 2,,5 2.5 1.9 2.0
Tiller number and leaf area.--T:le following results are evident:
the efect of deficiency in reducing both tiller number and leaf area
occurs in the order N > P > K; a decrease in potassium with high
levels of nitrogen and phosphorus leads to an increase in tiller num
ber, until the level of potassium is very much reduced; at a low phos
phorus level the efect of potassium defciency on tiller number is
completely masked, and, on leaf area, it is much less evident; nitrogen
defciency, in turn, masks completely the efects of phosphorus de
fciency; potassium defciency, unless very acute, has little efect on
meristematic activity, and, as is shown later, there is evidence that
potassium defciency does not inhibit protein synthesis.

Jo
0


9
w
.

r
N,KI-NsK,
oN,K,-N,Ks
7 N,K1 - NsKs
I I ~ _ _ _
Ol O2 O3 q 05 06 07 -`0 0' 0., O2 O Ol O 0.6 07
ToAl NItEN UPAKE lM UP' TOAL PCA$'11 UPAKE N<! P 0
FIG. I.-Relation between total yield and amounts of nitrogen and potassium
taken up. One nutrient in minimum means other Jutricnt in excess.
Yield.-The rClation of yield (total dry weight) to the amount
of nitrogen and potassium absorbed is shown in Figure 1. In the case
of nitrogen, both where nitrogen is minimal (N1Kl -NsKl) and
where nutrients are given
i
n a constant ratio (N lKl -N oK5)' the
relation of yield to nutrient absorbed is linear (law of minimum).
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MINERAL NUTRITION OF PLANTS 561
Yield plotted against potassium absorbed shows the same relation
when both nutrients are balanced (NIKl -NsKs), but when potas
sium is minimal (NIKl -N1Ks) the typical Mitscherlich relation
appears. In both cases, when yield is plotted against the amount ab
sorbed of the nutrient in excess, a curve of yield with increasing slope
is obtained. Chemical analysis of the plants elucidates these relations.
N,I,_
N,K2 N,K; ___ 0
N,K4 -.. _ N,Ks 0"".
N,K,
N"K, .. _0 NaK, _ ..
N4K,_ .. . Nsl'o ____ o
N,K,_
N2K,_-o N:K3_.-
N4K4___ _ Ns Ks 0--__ 0
2 ; 4 5 I 2 3 4 i I 2 3
4
FIG. 2.-Nitrogen content of barley plants sampled at fortnightly intervals.
On Jeft, series with potassium in minimum, nitrogen in excess; in center, nitro
gen in minimum, potassium in excess; on right, balanced series. (Abscissa
represents number of sample.)
For nitrogen they are shown in Figure 2. The series with nitrogen
in minimum (N,K, -NoK,), as well as the balanced series, shows
that in the early stages of growth large diferences in nitrogen con
tent are apparent, the higher internal concentration corresponding
with the higher external concentration. During the life cycle the in
ternal concentration falls and, eventually, in all series reaches almost
the same low level. This is in marked contrast to the relation in the
series with potassium in minimum and nitrogen in excess (N lKl -
NIK5)' Here, in the early stages, all the series show the same nitro
gen content in the tissues, but as development proceeds the series
receiving the least potassium now shows the highest nitrogen content.
Evidently the amount of growth made is the determining factor, the
larger the fnal size the lower the fnal internal concentration of nitro
gen. Similar relations hold for interactions of nitrogen and phos
phorus, as well as phosphorus and potassium. This explains fully
the so-called "luxury consumption" of nutrients present in excess.
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562 ANNUAL REVIEW OF BIOCHEMISTRY
The results will vary, however, as between nitrogen, phosphorus, and
potassium owing to their different relations to meristematie growth.
Where nitrogen is in excess, and .also to a less extent in the case of
phosphorus, growth occurs, but excess of potassium does not have
this efeLt. If, in Figure 1 at a given amount of nitrogen absorbed, a
line is drawn parallel to the ordinate, it is seen that the yield increases
with increase in potassium supply up to the "balanced" dosage, but
further increase in potassium lead: to no further increase in yield. If
the same is done for potassium uptake, the yield increases beyond the
balanced dose as nitrogen supply increases .. As shown in Figure 2 the
lower the level of potassium the higher the fnal concentration of
nitrogen when the supply of nitrogen is in excess of the balanced dose,
and this accumulation of nitrogen leads to the excessive tillering noted
above. Such plants have a very low assimilation rate, the stems fail
to elongate, and the ears although diferentiated fail to emerge. Such
plants are therefore completely sterile and the nitrogen taken up is
not efciently used. The Mitscherlich curve, therefore, expresses an
interaction efect between the nutrient in minimum and that in ex
cess: where nutrients are supplied in balanced proportions the law
of the minimum holds.
Balance of nutrients.-This question of salt balance is of very
obvious importance. One of the inherent difculties in evaluation of
the efects of any nutrient salt is that the ions are always introduced
in pairs, and therefore the combined efect of anion and cation is
always obtained. The method of three salt solutions introduced by
Shive sought to establish the most efective combination of six ions.
In order to evaluate the nutrient efect of single ions from such re
sults a statistical method alone is possible. Beckenbach, Wadleigh &
Shive (6) have published such an analysis for corn grown in a series
of six salt solutions. They establish the preponderant efect of NOs-,
and the almost negligible efect of P04 and S04=. The cations, K+
and CaH, gave increases in yield throughout the range studied. These
results are in complete agreement with a previous analysis of data
for potatoes reported by the reviewer (18).
Relation of growth to concentration of nutrients. -The early
stages in development are concerned mainly with the accumulation
of nutrients which proceeds rapidly in the tissues (Fig. 2). The rate
of uptake for all nutrients studied has been found to be proportional
to external concentration up to the stage at which foral diferentia
tion begins (six weeks in barley sown in May). The maximum rates
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MINERAL NUTRITION OF PLANTS 563
of uptake of nitrogen and phosphorus are shown in Figure 3. From
germination onwards the rate of uptake increases as the root system
expands. Two opposing processes regulate the rate; with expansion
of the root system the rate tends to rise and, as uptake increases, the
external concentration falls by depletion of the solution. As a result,
therefore, the rate of uptake reaches a maximum value early in the
life history. Since growth is an exponential process with an ever
increasing number of primordia laid down as new leaves and tillers,
0
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O
D Q

R
5300

M
Z

t VOO
w
3
t
C
' 1
W
!
e
INCREASINQ N A NsK,-N1K,
Do. NsK x NsKs-N,K,
Do. P N,Ps-N,P,
O-----
r
--------------
-

I Y9
FIG. 3.-Relation between rate of uptake of nutrients and external concen
tration; experiments with barley six weeks from germination. (External con
centrations of Nand P 205 are represented on the abscissa.)
the requirement of nutrients for further growth and maintenance
tends also to an exponential increase; meanwhile, the external solu
tion is depleted and, therefore, the rate of supply falls. The critical
point at which demand overtakes supply marks the maximum in rate
of uptake. This point coincides with the appearance of defciency
symptoms in the leaves at emergence, and has been called the stage of
"internal starvation" (10). Up to this stage leaf-area growth and
dry-weight increase are e7ponential (17). Where high levels of nu
trient are presented at the outset, or where nutrient supply is con
tinuously renewed, internal starvation supervenes nevertheless. This
has been shown in barley to be due to a sudden fall in the rate of up-
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56 ANNlAL REVIEW OF BIOCHEMISTRY
take by the roots. A similar cessation of uptake was found in cotton
by Crowther (10), and is related in both cases to the onset of the
reproductive phase.
The rate of development is proportional to the rate of uptake and
therefore to external concentration. For each concentration, there
fore, if not too high, the rate of uptake and rate of depletion are
proportional to external concentration. Maxima in tiller number and
leaf area should therefore occur at the same time independently of the
concentration at which the plant is grown, and this has been shown
to be the case with phosphorus and potassium in minimum.
The onset of internal starvation has an obvious bearing on the
optimal time at which nutrients should be supplied, and on recovery
from starvation. Tibeau (52) has reported on the time factor in the
utilisation of nutrients by hemp, and shows that recovery from pro
longed starvation of nitrogen is very slow. An interesting relation to
sex diferentiation is stressed, low nitrogen producing male fowers
and high nitrogen producing female flowers.
A comparative study of rye and oats in relation to mineral nutri
tion has been made by De1eano & Gotterbar (14). Losses of potas
sium, calcium, and nitrogen occur after fowering in rye, whereas oats
lose potassium only. The concentration of the sap is maintained
con
stant, when desiccation after fowering begins, by excretion of salts
by the roots.
Diferential m'ietal response.-Reference may here be made to
the efciency with which diferent varieties utilise nutrients in yield
production. This so-called "diferential varietal response" was frst
established with fve varieties of barley by Gregory & Crowther (19,
20). Lynes (27) has examined the phosphorus nutrition of twenty
one varieties of corn and has noted different symptoms of deficiency
and diferent rates of tilisation in the early stages. Large diferences
in susceptibility to phosphorus defciency were noted among the va
rieties. A high phosphorus requirement acted as a recessive in hy
brids. The response to phosphorus is correlated with the number and
character of the roots.
Lamb & Salter (26) established diferential response by statistical
analysis of yield with seventeen varieties of wheat grown at diferent
levels of fertility. Diferential response in wheat has been shown also
by Woodford & McCalla (58). Crowther and Crowther, Tomforde &
Mahmoud (11, 12) in large-scale field experiments have established
the same for va:rieties of cotton.
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MINERAL NUTRITION OF PLANTS 565
Symptoms of defciency.-There is general agreement as to the
symptoms of nitrogen and phosphorus starvation. The symptoms of
potassium starvation, however, have been variously described and are
discussed by Richards & Templeman (39). These may be either a
light yellow colour of leaves associated with succulence and very rapid
death of the leaves, or a dark green colour associated with white or
brown spots on the leaves. The frst type is characteristic of potassium
starvation with the standard manuring used in these experiments, in
which a high level of sodium is maintained. An additional efect under
such conditions of low potassium and high sodium is the continued
tillering which occurs after eight weeks, when, with ample supply of
potassium, tillering ceases. The frst crop of tillers dies and is suc
ceeded by a second and third cycle which in turn die without emer
gence of ears. In the second type described by American investigators
( see 39 for references) meristematic activity ceases and growth rate
falls. This second type of potassium defciency is characteristic of
conditions of high calcium and low phosphorus supply. Shih (49)
has shown that even in the presence of high calcium the frst type of
defciency symptoms is produced by adding extra sodium. Sodium,
therefore, has a specifc efect on growth. Scharrer & Schropp (44)
fnd no relation between sodium and the uptake of other nutrients,
but state that increasing concentration of sodium leads to greater up
take of potassium as well as sodium. Shih on the other hand fnds
that a decrease in potassium in all series studied leads to an increase
in sodium uptake.
Schneider (46) has examined the efects of defciencies on the
anatomical structure of Pelargonium, and the efects on the proto
plasmic structure of the leaves of Elodea are described by Kalch
hofer (25).
Water content.-Results in these experiments, discrepant from
those of American investigators, have been consistently obtained;
thus potassium defciency of the frst type (high sodium, low calcium)
is associated with increased succulence, and that of the second type
(high calcium, low phosphorus) with low water content. The work
of Shih (49) has shown that the water content with potassium def
ciency depends on the levels of sodium, calcium, and phosphorus. The
relation of water content to relative concentrations of sodium, cal
cium, and potassium, as found by chemical analyses of green leaves
in barley, are represented in Figure 4 in which the contours are of
equal water content (water/dry weight). The manurial combinations
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S6 ANNUAL REVIEW OF BIOCEMISTRY
used are indicated below the diagram. The following results have
been shown to be statistically ver signifcant: (a) Phosphorus content
has little efect on water content at a high potassium level, but an in
crease in phosphorus defciency reduces the water content as the potas
sium level falls, i.e., reduction of potassium increases the water content
A C high Na + CW C"
1 = medium u + medium Ca
C high Ca without Na
L 7 DW phosphorus
H = high phosphorus
K, K., K. " levels of potassium defciency
FI. 4.-Re1ation between water content ad relative proportions of cations
in green leaves 0: barley. Contours are of equal water content.
much more at high than at low phosphorus levels. ( b) Reduction in
water content due to a phosphors defciecy is much more evident at
high sodium than at high Calcium levels. (c) At a high sodium level,
potassium defciency leads to an increase in water content; in balanced
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MINERAL NUTRITION OF PLANTS 567
sodium-calcium series the efect of potassium is less marked; in a
high calcium series, reduction of the potassium level leads to reduc
tion in water content, but a further reduction in potassium leads to a
second increase. These results completely clear up the discrepancies
in the fndings of diferent investigators.
Nitrogen metabolism.-A detailed study of the various nitrogen
fractions in the tomato has been made by Clark (9) on plants fed
with nitrate and ammonia. The methods for study of nitrogen metabo
lism are dealt with by Orcutt & Wilson (34). The so-called reducase
activity of expressed sap has been reinvestigated by Sommer (50)
who fails to fnd evidence for catalytic reduction of nitrate to nitrite
in absence of light.
A study of defciencies in nitrogen, phosphorus, and potassium
(1/10 standard) on the nitrogen metabolism of barley leaves has been
published by Richards & Templeman (39). The successive leaves of
the main shoot at full emergence and during senescence were studied.
It should be noted that the number of successive leaves is almost in
dependent of manuring, ranging from ten to twelve in extreme cases.
The following fractions were estimated: total nitrogen, protein, total
amino, amide, nitrate, and residual crystalloid nitrogen. In general,
total nitrogen and most of the fractions reach a maximum in the
second to fourth leaf, decline to a minimum in the eighth or ninth
leaf and again rise in the last leaves. Two sources of nitrogen are
available: newly absorbed nitrogen and nitrogen liberated by prote
olysis. A study of the conditions leading to protein breakdown in
relation to chlorophyll destruction has been published by Michael (30).
Nitrogen defciency leads to low content in the leaves, but the frac
tions are present in the same proportions as in fully manured plants,
indicating a normal course of protein synthesis. Phosphorus def
ciency leads to reduction in protein content even at emergence, and a
rapid fall during senescence of individual leaves. The characteristic
efect is the accumulation of amide nitrogen and a less marked accumu
lation of aino nitrogen and nitrate in the later leaves. A check in
protein synthesis at the level of amide is indicated, and this failure of
protein synthesis leads in the case of phosphorus, as in nitrogen
defciency, to a low meristematic activity as shown by reduced tiller
production and individual leaf size and leaf area. Potassium defciency
(with high sodium, low calcium) is characterised by: (a) a marked
increase in amino and amide nitrogen, though they appear in normal
proportions; (b) accumulation of nitrate nitrogen in later leaves;
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568 ANNUAL REVIEW OF BIOCHEMISTRY
(c) a very rapid breakdown of protein during senescence of leaves. At
emergence the leaves show a normal protein content. It is held thqt po
tassium is not primarily associated with protein synthesis but is neces
sary for maintaining the protoplasmic complex, and in its absence
rapid proteolysis occurs. In consequence of this protein breakdown,
soluble nitrogen fractions accumulate throughout the plant. The ef
fects of high calcium and low sodium in modifying these conclusions
are not yet known.
Carbohydrate metabolism.-The eifect of nitrogen supply on sugar
prod uction in sugar cane has been studied by Das (13). High nitrogen
supply increases succulence, and the variations in water content are
related to diferent absorption of inorganic salts. Hydrtion is corre
lated with growth and the content of reducing sugar and sucrose. The
sucrose content of expressed sap is highest with nitrogen defciency,
but no relation to polysaccharides is found. It is concluded that the
inorganic constituents, acting on the enzymes are the chief controlling
fctors.
A study of defcienies in nitrogen, phosphorus, and potassium
0/10 standard) on the leaves of barley has been made by Gregory &
Baptiste (22) using successive leaves on the main shoot, both at
emergence and during senescence. The experiment was repeated in
two years with similar results., The standard manuring with high
sodium and low calcium was used.
Nitrogen deficiency has no consistent efect on reducing sugar,
while total carbohydrate is greatly increased. Phosphorus defciency
increases free reducing sugar, but total sugar is less afected. Potas
sium defciency lowers redu!ing sugar, and leads to a very low total
sugar content. No consistent diferences in sugar content appear in
the defcient series until after the emergence of the fourth leaf, at
which time tillering begins. A minimum sugar content occurs at this
time. Except in the potassiumdef\ient series the sugar level rises
from this point throughout the me.jor part of the life cycle. The
course of this ri:se depends on the mode of application of the manures.
Senescence in the earlier leaves leads to a fall in sugar content but
in the later leaves the opposite efect appears, the change-over occur
ring in diferent leaves in the various series. Potassium defciency
alone shows, in all cases, a consistent fall in sugar content during
:enescence. Characteristic diferences in the ratio of sucrose to free
reducing sugar are found: very high values in nitrogen defciency
very low in potssium defciency, and intermediate in phosphorus
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MINERA NUTRITION OF PLNTS 569
defciency. All the efects noted were highly signifcant statistically.
The high value of reducing sugar in the phosphorus-defcient leaves
is shown to be related to the high ratio of amide to total amino nitro
gen in this series,' and a sudden increase in both values occurs at the
time of emergence of the fourth leaf. The sugar level is related to
carbon assimilation, translocation, protein synthesis, and respiration.
High sugar content with nitrogen defciency is due to high assimila
tion rate, low respiration, low protein synthesis, and low meristematic
activity. Lower assimilation rate with phosphorus defciency is ofset
by the low meristematic activity so that the sugar content does not
depart much from normal. In the potassium-defcient plant the very
low assimilation, high respiration, active protein synthesis, and exces
sive meristematic activity all contribute to a very low sugar content.
The investigations of the efect of potassium defciency on the
barley leaf have been extended to cover conditions of high calcium
and low sodium. Three levels of potassium (Kl' Kg, K5) combined
. with two levels of phosphorus have been used. A fructose anhydride
has recently been isolated from water extracts of barley leaves by
Archbold & Barter (1) and estimates of this carbohydrate fraction
were included in this experiment.
Some of the results obtained are entered in Table V. A key to the
manurial combinations is given. The values entered are means for
the ten successive leaves, expressed as equivalent hexose in milligrams
per gram of leaves (fresh weight).
The following results appear: (a) The total carbohydrate content
is greater, on the average, in the series with high calcium than with
high sodium; (b) potassium defciency leads to a great reduction in
all fractions in the presence of high sodium, but in the series with
high calcium reduction in carbohydrate is not at all evident at Ks level
and is only very small at K. level; (c) reduced phosphorus in the high
sodium series alone leads to an accumulation of reducing sugar and
fructosan. Fructosan accumulation is also dependent on the potassium
level, low potassium leading to a fall in fructose cotent.
Efect of nutrient defcienc
y
on rpirtion.-The literature on the
efect of nutrient defciency on respiration and assimilation is very
limited. Much of the previous work sufers from the following de
fects: (a) insufcient regard has been paid to the changes in rate
which take place during development; (b) evaluation of statistical/
signifcance is seldom attempted.
Estimations have been made of respiration, etc., of successive
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570 ANNUAL REVIEW OF BIOCHEMISTRY
leaves on the main stem at the time of complete emergence by Gregory
& Richards (21), and Richards (37). The use of observations made
on all leaves simultaneously has been avoided for the reasons given
by Richards (38) who points out that this method confounds the
diferences of metabolic rate in leaves formed at diferent stages in
the life history of the plant with the senescent changes in individual
leaves; the metabolic history of each leaf varies and due egad must
b paid to this fact. Only results confrmed by statistical analysis
are accepted.
TABLE V
CARBOHYDRATE FRACTIONS AS EQUIVALENT HEXOSE
(mg. per gm. fresh weight)
Reducing
Sugar Sucrose Fructosan
HAKl ..... . .... ..... 2.4 16.3 5.9
HAKs ... . . .......... 2.7 14.9 1.7
HAKs .......... . .... 1.9 7.3 1.1
LA . . . . . ........ .. 4.3 16.7 11.3
LAKs .... . .... " ." . , 3.5 16.4 4.4
LAK
G
. .... . .... . . ... 2.8 8.4 3.7
HCK1 ....... . . ... . .. 4.5 17.3 5.4
HCKa .. . . .. . . ... . . .. . 4. 7 16.4 3.4
HCK5 .. . . . . . . . . . .... 4.1 18.9 2.6
LCK1 0 0 0 4.6 18.5 9.6
LCKs ........ . ...... 4.6 19.9 9.3
LCK5 .. . ...... . ..... 5.3 15.4 5.2
H 7 high phosphorus P = high sodium
= low phosphorus (lIS) C = high calcium
K,. K
.
, K. = levels of potassium (I, 1/9, 1/81)
Total
Sugar
24.7
19.4
10.2
32.3
24.3
15.0
27.2
24.5
25.8
32.6
33.9
26.0
The relation of respiration to water content is discussed by
Schlieper (45).
Nitrogen defciency in barley always leads to a reduction in the
rate of respiration (21, 48). This agrees with the fndings of MUl
ler & Larsen (3:2), Ruhland & Ullrich (41), and Hamner (23).
The respiration of successive leaves within a series shows an in
crease at frst and reaches a maximum at the second to foLrth leaf;
it declines to a minimum at the eighth or ninth leaf and rises subse
quently. This agrees with the changes in total nitrogen already men-
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MINERAL NUTRITION OF PLANTS 571
tioned (39). Phosphorus defciency at the level 1/5 results in no
signifcant reduction in respiration (21), but on further reduction of
the phosphorus supply a progressive decrease in respiration results.
Jones (24) has investigated the efect of phosphorus supply on
etiolated wheat seedlings and has shown a small increase in carbon
dioxide production with addition of phosphorus. He concludes that
phosphorus acts by its association with carbohydrate metabolism ad
by its regulation of protein synthesis.
Potassium defciency at the level Ka (1/9) leads to a very great
increase in respiration but further reduction in potassium leads to a
fall in carbon dioxide production, which in extreme starvation is below
normal (37). These results were obtained with high sodium and low
calcium, i.e., pale green succulent leaves.
Respiration of leaves under conditions of high calcium, i.e., dark
green leaves with low water content has since been studied. Again
great increases in respiration are found with potassium defciency;
the mean respiration rate of successive leaves in the high calcium
series at two levels of phosphorus are given in Table VI.
TABLE VI
RESPIRATION RATE; MEAN OF ALL SUCCESSIVE LEAVES
(mg. CO2 per gm. dry weight per hour)
HCK, ............. 4.38 LCK, ............. 4.70
HCKa ............. 5.42 LCK3 ............. 4.60
HCK5 ............. 6.51 LC ...... .. . .... 4.9
the increase in respiration with potassium defciency is thus found
to be general in barley. The efect of potassium is dependent, how
ever, on the level of phosphorus; the efect of potassium defciency is
masked when phosphorus is kept at a low level.
The relation of respiration to carbohydrate and nitrogen metabo
lism has also been investigated. Sen (48) studied the relations in
series of minimal nitrogen and potassium
Estimations of protein, amino nitrogen, reducing sugar, and total car
bohydrate were made. In all series with the single exception of
N lK
5 signifcant high positive correlations between respiration and
protein content were found. As also observed previously (39) potas
sium defciency gave a very high amino nitrogen content, and in this
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57 ANNUAL REVIEW OF BIOCHEMISTRY
series ( N1Kl. - N1K5) a signifcant high correlation of amino nitro
gen and respiration was noted. Very high values of sugar in thC
nitrogen-defcient and very low values in the potassium-defcient
series were found (d. 22). Thus the nitrogen-defcient series had a
low respiration associated with high sugar content and the potassium
defcient series, fhe converse. Indeed in no case within a manurial
series was there a signifcant direct correlation between respiration
and either total or reducing sugar content with the single exception of
the N1K5 series which gave a high positive value with total sugar. The
partial correlation, after eliminating amino and protein nitrogen ef
fects, was still very high and signifcant. and in the NiKa series also
a signifcant positive partial Lorrelation of resp
i
ration rate and sugar
was obtained when the efect of amino nitrogen was eliminated. It
thus appears, as Richards surmised (37), that with very great potas
sium defciency carbohydrate becomes the controlling factor in respi
ration, but in no other case is this so. The very high correlations of
respiration with protin (nitrogen defcient ) and amino nitrogen
(ptassium defcient) are unafected by eliminating the efect of sugar
content of the leaf.
A similar study has been madL of respiration in the series with
minimal phosphorus and also in interaction with potassium (balanced
series) in relation to carbohydrate and nitrogen fractions. When
potass
i
um
i
s reduced at the lower level of phosphorus from PaKi to
PaKa there is again a sigifcant increase in respirati on but at the
P g level reduction in potassium has no efect. A similar relation to
amino nitrogen has been obtained. A striking similarity is thus .seen
throughout between the efects of potassium defciency on respiration
and amino nitrogen content. Schwabe (47) has obtained striking ef
fects of amino acids on the oxygen. uptake of Elodea leaves. As a
result of feeding with tyrosine and glutamic acid, increases of 30 per
cent in respiration wCre obtained. By periodical removal from amino
acid solution to water the efect di sappearedy to reappear again on
replacing in the amino acid solution. The amino acid was thus used
up but the respiratory quotient was unafected, indicating that the
amino acid is not direLtly oxidised. It is suggested that the amino
acids stimulate generHl oxidation by acting as hydrogen acceptors.
Returing to the experiments with barley it thus appears that the
efects of phos
p
horus defciency on amino nitrogen accumulation and
respiration are Oppos{!d, a decrease in respiration rat being accom
panied by an increase in amino nitrogen; potassium starvation, on
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MINERAL NUTRITION OF PLANTS 573
the other hand, leads to an increase in amino nitrogen as well as in
respiration. In the series Pg and P5 the correlations of respiration
with protein are not signifcant, but high positive correlations with
reducing sugars are found. The efects of phosphorus and potassium
defciencies are thus very diferent, the efect of the former apparently
working through carbohydrate relations, the latter through nitro
gen metabolism.
Feeding experiments.-A brief reference may here be made to
experiments performed to elucidate the relations mentioned above by
feeding leaves from starved plants of barley with the particular nutri
ent in defciency. Said (42) worked with potassium-starved leaves
and showed that feeding with potassium alone had no efect, which
demonstrates that the efect of potassium is indirect. H
o
wever, feed
ing with sugar (glucose, fructose, or sucrose) increased the respira
tion rate.
With phosphorus-starved leaves feeding experiments have been
performed by Sankaran (43). Leaves removed at diferent stages
during the life cycle were fed as follows : (a) distilled water, (b) su
crose in excess, (c) sucrose and sodium phosphate, and (d) sucrose,
sodium phosphate, and ammonium nitrate. In addition to carbon
dioxide production, estimations of phosphorus content, protein nitro
gen, amino nitrogen, amide nitrogen, and residual nitrogen were made
before and after the period of respiration (three or four days) .
It was found that respiration was increased very considerably by
addition of phosphorus and sugar, and an additional increase was
obtained by giving ammonium nitrate with the sugar. The magnitude
of the increase, however, depended upon the stage at which leaves
were removed, the efect being greatest in the earlier leaves; thus the
efect of feeding with phosphorus was greatest when the leaves nor
mally contained most phosphorus. In fact the efect of phosphate
addition on increasing the respiration is greatest when the respiration
is normally highest. Schwabe (47) points out a similar efect of
amino acids on Elodea.
It was also observed that considerable protein hydrolysis occurred
during respiration but this was always less in the leaves fed with
ammonium nitrate.
The conclusion reached from these experiments, though the evi
dence cannot be presented here, is that the main efect of phosphorus
on respiration depends on its relation to nitrogen metabolism, and the
efect on carbohydrate metabolism is secondary. The evidence is not
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574 AmAL REVIEW OF BIOCHEMISTRY
inconsistent with the view that carbohydrate is not oxidised unless it
is frst drawn into the cycle of nitrogen metabolism. On this view the
efect of potassium defciency is related to amino nitrogen content
because the level of this fraction measures the rate of proteolysis. As
at the same time, as has been shown, the protein content of potas
sium-starved leaves is at frst very high, it follows that protein syn
thesis as well as hydrolysis must be taking place in potassium-starved
leaves very rapidly; this, it is suggested, is related to the high respira
tion rate. Further discussion cannot be pursued here.
White (55) , working with Lemna, fnds during recovery from
nitrogen starvation that the carbohydrate which has accumulated
during starvation is very rapidly oxidised. The respiration rate may
reach a value of fve times that previously shown under nitrogen de
fciency. Rohde (4) also discusses the efect of potassium on respira
tion. He suggests that potassium regulates the distribution of iron
in the plant ; the iron, in turn, acts as a catalyst for oxygen uptake.
The increased oxygen supply inhibits anaerobic respiration, and the
. efect of potassium therefore is related to the relative rates of aerobic
and anaerobic respiration.
Efect of nut1ient defciency on carbon assimilation.-Experiments
have been performed in which the assimilation rates of barley leaves
' from plants deficient in nitrogen, phosphorus, and potassium have
been studied in full daylight, using air as the source of carbon dioxide.
Successive leaves were again used, while attached to the plant. The
air supply was very rapid, the assimilation chamber shallow, and the
leaf area small so that ma. ximal rates of assimilation were obtained .
. Nitrogen defciency at the levels N3 ( l/9) and N5 ( 1/81 ) were studied
by Chinoy (8) . No reduction in assimilation rate was found as may
-be seen from the following values :
Control, fully manure . . . . . . . . . . . . N
l
Medium defciency . . . . . . . . . . . . . . N
a
Control, fully manured . . . . . . . . . . .
Nl
Extreme defciency . . . . . . . . . . . . .
N
5
(
1
/9)
Assimilation of Carbon Dioxide
(mg. per SQ deem. per hour)
19. 4
J
Mean value of 17
20. 1 determi nations
Mean value of 11
(1/81)
2. 1 1
20. 1 determinations
This confrms the previous fndings of Gregory & Richards ( 21 ) .
Similar experiments with phosphorus and potassium-defcient leaves
show, after the fourth leaf, a rapid fall in assimilation which reaches
a minimum in the eighth leaf and subsequently rises ( cf. 21 ) .
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MINERAL NUTRITIO
N
O PLANTS
575
The results with nitrogen defciency are not in agreement with
those of Muller (31 ) , Muller & Larsen (32) , and Hamner (23) ,
who record a reduction i n assimilation rate due to nitrogen defciency.
This discrepancy is in part due to the methods used by these investi
gators : no efort was made to separate the senescent efect of lack of
nitrogen on individual leaves from the efect on the leaf at the time
of its emergence. There is no doubt that nitrogen defciency leads to
a more rapid senescence of leaves and this is even more evident with
phosphorus and potassium defciencies. Unless these factors are sep
arated discordant results must be expected. Gassner & Goeze ( 1 5, 1 6)
have stressed the importance of the interactions of nitrogen and po
tassium on assimilation rate. In their experiments the frst leaf alone
was used, which, as Steward (51 ) has rightly pointed out, would be
least likely to sufer from nutrient defciency owing to stores in the
seed. In our experiments the marked efects on leaves just emerged
appear at the ffth leaf, and it is interesting to note that the frst four
leaves of barley are preformed in the embryo, the ffth being difer
entiated during germination. It would thus appear that the conditions
under which the primordia are laid down are very important in deter
mining the reaction to nutrient supply. In the experiments of Gass
ner & Goeze the efects on assimilation in the diferent series become
more marked as the experiments proceed ( 16) , and indeed, with leaves
ten days old, changes in the nitrogen supply ( 1/10, 1, and 5) were
without efect on the assimilation rate. Senescence was, therefore,
primarily investigated in this work. The relation of potassium to dry
matter accumulation is discussed by Maiwald & Frank (28) .
The results obtained with barley by direct estimations of assimila
tion are in agreement with estimates of net assimilation derived from
sampling data. The method is described by Gregory ( 17) ; it is, with
out doubt, the best method available for studying average assimilation
rates over long periods. Data from such estimates are given by Greg
ory & Baptiste (22) . By this method nitrogen defciency up to maxi
mal leaf area (i.e., until senescence of leaves begins ) shows no efect
L assimilation [Gregory ( 1 7) , Mathur (29) ] . This has been shown
for cotton also by Crowther ( 10) . Verma (53) showed by the same
method with barley that phosphorus and potassium defciencies re
duce assimilation in the order stated. Watson (54), from sugar
analyses on potato, concludes that addition of potassium chloride in
creases assimilation rate. This important paper, however, is primarily
concerned with diurnal variations in carbohydrate fractions and in
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576 ANNUAL REVIEW OF BIOCHEMISTRY
water content. Vhite ( 55), working with Lemna, found no reduc
tion in net assimilation as a const!quence of nitrogen starvation,
though low respiration and low multipli ation rate were symptomatic.
Further, the same investigator fnds a low assimilation rate to be
characteristic of potassium defcienLy ( 56) . During the course of
potassium starvation dry matter accumulates in the fronds so that the
dry weight per unit area reaches a high constant level. On renewing
the potassium supply the accumulated dry matter rapidly disappears.
Enzyme extracts show a reduced capacity for starch hydrolysis with
potassium defdency+ The main efect of potassium is attributed to
the regulation of carbohydrte meta.bolism by control of the starch
sugar balance.
In conclusion, to indicate some of the complexities of the relations
of assimilation to nutrient ions, reference should be made to, the im
portant papers of Arens (2, 3, 4) who has investigated the assimila
tion of aquatic plants and has shown that calcium bicarbonate mole
cules are absorbed at the lower surface of the leaf while calcium
hydroxide escapes at the upper. This polar transport is found only
in the light, no a.bsorption occurring in the dark. In the case of potas
sium bicarbonate absorptiong potassium carbonate escapes from th(
upper surface.
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MINERAL NUTRITION OF PLANTS 577
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INSTITUTE OF PLANT PHYSIOLOGY
hlPERIAL COLLEGE OF SCIENCE AND TECHNOLOGY
LONDON, ENGLAND
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