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BENEFICIAL EFFECT OF CREATININE AND CREATINE
ON GROWTH'
J. J. SKINNER
(WITH ONE FIGURE)

This paper embodies a seriesof experimentson the influenceof

creatinine and creatine on seedling wheat. These experiments
were made in an endeavor to throwlight on the action of organic
manures in soils, and the influenceof soil organic matter on pro-
ductivity. Creatininehas been discoveredas a soil constituentin
this laboratoryby Dr. E. C. SHOREY,2 and an account of its occur-
rence and propertieswill be given elsewhere. This nitrogenous
constituentoccurs plentifullyin animal products,wine, meat, etc.,
but has recentlybeen found in these laboratories by Dr. M. X.
SULLIVAN3 to be a constituentpart of many plants and seeds,
and to occur in the medium in which plants have grown. The
general methods for studyingthe effectof creatinineon plants in
solution culturesis the same as that employedin connectionwith
the harmful soil constituent, dihydroxystearicacid, previously
reportedin this journal,4
Effectof creatinineon growth
Two sets of cultures,composedof the fertilizer
salts calciumacid
phosphate,sodium nitrate,and potassium sulphate in varyingpro-
portions,used singly and in combinationsof two and three,were
prepared,the proportionsvaryingin io per cent stages, thus mak-
ing a total of 66 culture solutions according to the plan in the
I Published by permissionof the Secretaryof Agriculture, fromthe Laboratory
of Soil FertilityInvestigations.
2 SHOREY,EDMUND C., The isolation of creatinine from soils. Jour. Amer.
Chem. Soc. 34:99. I9I2.
3 SULLIVAN, M. X., The origin of creatinine in soils. Jour. Amer. Chem.
Soc. 33:2035. I91I.
4 SCHREINER, O., and SKINNER, J. J., Some effectsof a harmfulorganicsoil con-
stituent. BOT. GAZ. 50: i6i. i9io; Ratio of phosphate,nitrate,and potassium on
absorptionand growth. BOT. GAZ.50: I. I9IO.
BotanicalGazette,vol. 541 [I52
I9I2] SKINNER-CREATININE AND CREATINE

papers cited. Young wheat seedlingswere grownin this series of

solutionsfromMarch 3 to March I5. To one set of the 66 cultures
only the nutrientsalts were added, to the second set 50 ppm. of
creatinine were added to each culture. Every three days the
solutionswere changed and analyzed.
When the two sets of cultures had grownfor several days, it
was noticeable that the creatinineplants were better developed,
having broader leaves and longerand well developed roots. This
was morenoticeablein some of the fertilizer mixturesthanin others.
The total growthmade in the 66 culturesof nutrientsalts with-
out creatinine,designated as normal cultures,was i66. 7 grams as
against i8i . 2 gramsin the case of the 66 cultureswith 50 ppm. of
creatinine. Putting the normalat ioo, the latter becomes IO9, or
an increase of 9 per cent as an average of the 66 cultures. As
already mentioned,the effectwas much more pronouncedin cer-
tain fertilizercombinations,especially those containingno nitrates,
or those low in nitrates. The effectsof creatininein these cultures
will now be consideredin detail.

Effectof creatinine on growth in cultures containing no nitrate

Table I gives the growthof two sets of cultures composed of
mixturesof phosphate and potash, varyingin io per cent stages,
TABLE I
SHOWING THE EFFECT OF CREATININE ON GROWTH IN CULTURES CONTAINING NO
NITRATES

PP)M. OF FERTILIZER INGREDIENT IN CULTURE GREEN WEIGHT OF CULTURE

SOLUTION IN GRAMS
No.
P"0s NH3 K0 Without With
creatinine creatinine
I . . . . . . . . .. 0 0 80 I.400 I*576
2. . . . .... . 8 0 72 1.470 2.200
3.1. -.....I6
- . 0- -
64 1.950 2.IOO
4 - -......... 24 0 56 I.527 2.000
5 *. * . . . . . 32 0 48 1.490 2.200
6 ......... 40 0 40 I.558 2.408
7 - -. ......... 48 0 32 1.795 2-328
8. . . . . . . ... 56 0 24 I.540 2.400
9. . . . . .. ... . 64 0 i6 1.444 2.220
I 0.......... 72 0 8 I.400 2.I00
I I . ...... 80 0 0 I.I00 I.I50
I54 BOTANICAL GAZETTE [AUGUST

there being no nitrate in the solutions; the concentration was

8o ppm. of P205+K20 in each culture. To one set of cultureswas
added 50 ppm. of creatinine. In the fifthcolumn are given the
green weights of the cultures without creatinine,and in the last
column are given the weightsof the cultureswith creatinine. It
is apparent from these figuresthat the creatininehas caused a

.. .....
Is ~~~~~~~~~

.s* We sj 'X 4' 4 lb; # P

FIG. i.-Wheat plants growingin culture solutionscontainingvarious propor-

tionsof potash and phosphate(withno nitrates)without(a) and with (b) creatinine.

considerableincrease in growth. This is true in each of the iI

cultures. The total growth of the eleven cultures, without
creatinine, was i6 .674 grams against 22 .682 grams for the
cultures with creatinine. This is an increase of 36 per cent in
the creatininecultures.
The effectof creatininein cultureswith no nitrogenare shown
in the plants in fig. i. Cultures marked with the same number,
forinstance ia and ib, have similarfertilizerratios. The cultures
marked a have no creatinine,the numberswith the letter b have
I9I2] SKINNER-CREATININE AND CREA TINE I55

50 ppm. of creatinine. As shown in the photograph,the plants in

each culture containingcreatinine,regardlessof the proportionof
potash and phosphate,is largerthan the plants grownin a similar
solution without the creatinine. The increased growthis notice-
able in the roots as well as the tops. The tops in each case are
broader and taller, the roots are largerand betterbranched.

Effectof creatininein cultures containing8 ppm. NH3 as nitrate

Since creatininewas very beneficialin cultures containingno
nitrate,it is interestingto observe its effectin cultureswhich con-
tain a small amount of nitrate. Table II gives the result of the

TABLE II
SHOWING THE EFFECT OF CREATININE ON GROWTH IN CULTURE SOLUTIONS COMPOSED
OF FERTILIZER MIXTURES, CONTAINING 8 PPM. OF NH3 AS NITRATE

PPM. OF FERTILIZER INGREDIENTSIN CULTURE GREEN WEIGHT OF CULTURE

SOLUTION IN GRAMS
No. . _ -

P0os NH3 K.0 Without With

creatinine creatinine

I. . . . . . 0 8 72 I. 820 2. I90
2 . 8 8 64 2 470 3 100
3 .I6 8 56 2 748 3.250
4 **. . 24 8 48 2.907 3.420
5.. . 32 8 40 2.670 2.450
6 . 40 8 32 2.928 3.258
7. 48 8 24 2.526 3.340
8 .56 8 i6 2.600 3.000
9. .
64 .8 8 2.048 2.359
Io .......... 72 8 0 I.354 I*750

effectof creatinine on growth in culture solutions composed of

8 ppm. of NH3 as nitrate,and varyingamounts of phosphate and
potash, the total concentrationof each solution being 8o ppm. of
P205+NH3+K20. By comparing the figuresit is seen that the
growthwith creatinine,given in the last column,is largerthan the
growthwithoutcreatinine,given in the fifthcolumn. The differ-
ence, however,is not nearly so large as in solutionscontainingno
nitrate,presentedin table I. The total greenweightofthe cultures
composedof fertilizermixturescontaining8 ppm. of nitrogenwith-
was 24.07I
out creatinine gramsagainst28. I 17 gramsin the cul-
I56 BOTANICAL GAZETTE [AUGUST

tureswith creatinine,an increase of I7 per cent. In the cultures

with no nitratecreatinineproduced an increase of 36 per cent.

Effectof creatininein cultures with larger amounts of nitrate

It has been shownthat creatininewas verybeneficialin cultures
whichcontainedno nitrates. In a group of cultures,composed of
mixturesofphosphateand potash in different proportions,creatinine
increased the growth36 per cent. It has also been pointed out
that the beneficialeffectof creatininewas not so great in cultures
containing a small amount of nitrate. In a second group of
cultures,composed of mixturesof potash, phosphate, and 8 ppm.
of NH3 as nitrate,creatinineincreased the growthonly I7 per cent.
In table III are given the resultsof growthin cultureswith and
without creatinine,composed of mixtures of phosphate, potash,
and nitrogenhaving i6ppm. NH3 as nitrate. The green weights
of the creatinineculturesgiven in the last column of the table are
slightlylarger than the normal cultures, as shown in the fifth
column. The total greenweightof the cultureswithoutcreatinine
was 25. 5i6 gramsagainst 27. 573 gramsfor the cultureswith
creatinine,an increase of 8 per cent.
TABLE III
SHOWING THE EFFECT OF CREATININE IN CULTURES CONTAINING i6 PPM. OF NH3
AS NITRATE

PPM. OF FERTILIZER INGREDIENTS IN CULTURE GREEN WEIGHT OF CULTURES

SOLUTION IN GRAMS
No.
No. P205 NH3 K2. Without With
P~~O5 NH3 K~~~~O creatinine creatinine

0 i6 64 2.200 2.570
2.. 8 . i6
...... 56 ....
3.200 3.720
3-***........... i6 i6 48 3.500 3-500
4-***............ 24 i6 40 3.097 3.702
5- * *.........
* 32 i6 32 3.250 3.250
6... 40 . i6
...... 24 ...3.228 3-300
7 .- ** ........ 48 i6 I6 2.975 3.240
8... 56 . ..i6 . ..8 2.626
.... 2 -55I
9.......... 64 i6 0 I 1.440 1.740

In other cultures composed of the three fertilizeringredients

P205, NH3, and K20, but containing 24 ppm. of NH3 as nitrate,
creatinineincreased growthonly 2 per cent. Its effectin cultures
I912] SKINNER-CREATININE AND CREA TINE I57

composed of fertilizermixtures having more than 24 ppm. of

nitratewas uncertain; in some cases therewas a slightincreasein
growthand in otherstherewas a slightdecrease,that is, the growth
with thesehigheramounts of nitratein the solutionwas practically
the same in the normal and creatininecultures.
Before discussing furtherthe effectof creatinine,it will be
necessaryto recall the effectwhich nitrateshave on the growthof
plants in mixturesof the othertwo fertilizeringredientspotash and
phosphate. In work previouslypublished,5it was shown that the
better growthoccurredin the normal cultureswhen the threefer-
tilizerelementsP205, NH3, and K20 were present. It was best in
mixtureswhich contained approximatelyequal amounts of NH3
and K20 and a small amount of P205 (about i6 ppm.). The
growthin the culturescontainingthe threeconstituentswas much
greater than in the cultures containing only two constituents.
This was especially marked when nitrogenwas not in the compo-
sition. In illustrationof this, the average growthof a numberof
cultures,composedofmixturesofphosphateand potash in amounts
of 8oppm. of P205+K20, was i.ooo gram against 3.I55 grams as
the average growthof culturescomposed of mixturesof these two
ingredients,with an addition of only 8 ppm. of NH3 as nitrate,the
total concentrationof nutrients being the same. In a second
experimentconducted in a similarmanner,but at a later date, the
average growthof the cultures,composed of mixturesof phosphate
and potash, was o. 878 gram, and the average growthof cultures,
in mixturesof the three ingredients,containing8 ppm. of NH3 as
nitrate,was 2. I07 grams.
In the present experimentthe growthin the normal cultures
composed of varyingproportionsof phosphate and potash, com-
pared with the growthin mixturesof these two ingredients,with
8 ppm. of NH3 as nitrate added, is given in table IV. By a close
examinationof the figuresin this table, it is seen that the growth
in the mixturesof phosphate and potash is smallerthan in cultures
composed of mixturesof the three ingredients,though containing

5 SCHREINER, O., and SKINNER,J.J.,Ratio of phosphate,nitrate,and potassium

on absorptionand growth. BOT. GAZ. 50: I. I9IO. Some effectsof a harmful organic
soil constituent. Bull. 70, Bureau of Soils, U. S. Dept. Agric. 19io.
158 BOTANICAL GAZETTE [AUGUST

but 8 ppm. of NH3. The average growthof the cultureswithout

nitrogenis I .5I6 grams against 2.407 grams with 8 ppm. of NH3
in the fertilizermixture. Putting the growthof the cultureswith-
out nitrogen ioo, therelativegrowthof the cultureswithnitrogen
at
becomesI59, or an increaseof 59 per cent.

TABLE IV
SHOWING THE GROWTH OF CULTURES, COMPOSED OF FERTILIZER MIXTURES CONTAINING
NO NITRATE, AND 8 PPM. OF NH3 AS NITRATE, WITHOUT AND WITH CREATININE

PPM. OF FERTILIZER INGREDIENTS GREEN WEIGHT OF CULTURES IN GRAMS

IN CULTURE SOLUTION

No. Without creatinine With creatinine

P2O NH3 K20

No nitrate 8 ppm. NH3 No nitrate 8 ppm. NH3

I ...... 0 8o I-.400 ..... I*576 .....

2 0..... 8 72 ..... I.820 ..... 2.I90
3 .... . 8 0 72 I*470 ..... 2.200 .....
4 ..... 8 8 64 ..... 2.470 3.I00
5 ...... i6 0 64 I*950 ..... 2.IOO .....
6 ..... i6 8 56 .... 2.748 3.250
7...... 24 0 56 I*527 ..... 2.000 .....
8 .. 24 8 48 ..... 2.907 ..... 3.420
9 ...... 32 0 48 I*490 ..... 2.200 ...
10 ...... 32 8 -40 ..... 2.670 ..... 2.450
II ...... 40 0 40 *558 . 2.408 .....
12 ...... 40 8 32 2.928 ..... 3.258
13 . 48 0 32 I.795 ..... 2.328 .....
I4 ...... 48 8 24 2.526 ..... 3.340
I5 ...... 56 0 24 I*540 ..... 2.400 .
i6 ... ... 56 8 i6 ..... 2.600 .... I 3.000
17 ... 64 0 i6 I.444 ..... 2.220
I8. . ..... 64 8 8 ..... 2.048 ..... 2-359
19 ...... 72 0 8 I-400 ..... 2.IOO .....
20 ...... 72 8 0 . .... 1*354 I.750
2I ...... 8o 0 0 I.I00 ..... I.I50 .....

With 5o ppm. of creatinine in the solution, the cultures con-

tainingno nitrogenproduced bettergrowththan the corresponding
cultureswithout creatinine,as shown in the last two columns of
table IV. The differencebetween the last two columns is not as
markedin the creatinineset as in the corresponding columnsforthe
normalset. The average growthof the creatinine cultureswithout
nitrateis 2 .062 grams against 2.8I2 gramsforthe cultureshaving
8 ppm. of NH3 as nitratein the fertilizermixture. If the growth
of the cultureswithoutnitrateis put at ioo, the growthwith 8 ppm.
I9I2] SKINNER-CREATININE AND CREATINE

of NH3 in the fertilizermixturebecomes I36, or an increase of only

36 per cent. In otherwords,in the absence of creatininefromthe
cultures,the nitrate (8 ppm.) caused an average increase of 59 per
cent in the various cultures; in the presence of the creatinine
(go ppm.) the nitrate (8 ppm.) caused an average increaseof only
36 per cent. It appears, therefore,that plants supplied with
creatininedo not respondso markedlyto added nitrate,thus seem-
ing to indicate that the plant can utilize thisnitrogenouscompound
forplant syntheses.

Effectof creatinineon absorptionof fertilizersalts

The foregoingdiscussion has shown clearly the influenceof
creatinineon growthand its effectin culturescontainingno nitrates.
There remains to be discussed the effectof the creatinineon the
removal of nutrients from the solution during the growth of
the plant.
Mention has been made already of the fact that the concentra-
tion differencesproducedby the growthof the plants in the various
cultureswere determinedby making an analysis fornitratesat the
terminationof every three-daychange, and of the phosphates and
potassium on a composite of the solutions fromthe four changes.
It is thus possible to compare the results obtained under the so-
called normal conditions without the creatinine and under the
conditionswhere50 ppm. of creatininewerepresentin the solution.
The sum total of P205, NH3, and K20 removedfromsolutionby
the growingplants in the cultures containing all three of these
constituentswas i684 milligramsunder the normalconditions,and
I584 milligramsin the creatinineset. The figuresshow the total
of plant nutrientsto be slightlyless in the creatinineset, although
thegreenweightin thisset was 9 per centgreaterthan in the normal
set. The examination of the results for the three constituents
separatelyas givenbelow shows that thephosphateand potash were
slightlygreaterthan normal,as is demanded by the largergrowth,
whereasthe nitrateis considerablyless than in the normal set.
Phosphate.-The amount of phosphate stated as P205 removed
fromthe total numberof solutions duringthe experimentwas 364
milligramsforthe normal culturesand 383 milligramsfor the cul-
i6o BOTANICAL GAZETTE [AUGUST

tures containingcreatinine,a differenceof i 9 milligramsin favor

of the creatininecultures.
Potassium.-The amount of potash stated as K20 removed by
the plants in the total number of cultures was 760 milligramsin
the case of the normalculturesand 778 milligramsforthe cultures
with creatinine. As with the phosphate, the creatinine cultures
removeda littlemorepotash than the normalcultures,therebeing
a difference of i8 milligramsin favor of the creatinineset.
Nitrogen.-The total amountofnitrogenstated as NH3 removed
fromthe total numberof solutionsduringthe course of the experi-
mentwas 560 milligramsforthe normalculturesand 423 milligrams
forthe creatininecultures. The creatinineculturesthoughmaking
a largergrowthused I37 milligramsless nitrate.

Effectof creatine on growth

Creatine is closely related chemically to creatinine,the latter
being the anhydride of creatine. Both probably occur in soils,
manures,and greencrops, a discussionof whichis given in the two
other papers referredto. Experimentsin nutrientcultures with
creatinehave been conductedsimilarto thosewith creatinine.
The plants grewfromApril 22 to May 4. Afterthe plants had
grownforseveral days, it was apparent that the effectof creatine
was very similar to that of creatinine. The leaves were broader,
and furtherdeveloped than those of the normalculture. The roots
were longerand betterbranched. The plants growingin cultures
withcreatine,whichcontainedphosphateand potash but no nitrate,
were a great deal larger than similar cultures without creatine.
Like the creatinine,when small amounts of nitratewerein the fer-
tilizermixture,the beneficialeffectof creatinewas not so marked,
and in the presence of larger amounts of nitrate creatine had no
additional effects.
The total green weight of 66 cultures containingthe fertilizer
salts only, that is the normal set, was I74.4 grams,against i86.8
gramsforthe 66 culturescontaining50 ppm. of creatinein addition
to the fertilizersalts. This is an increase forthe creatinecultures
of 8 per cent over the normal cultures.
Table V shows the effectof creatine on growthin a numberof
I9I2] SKINNER-CREA TININE AND CREA TINE

culturescontainingvaryingamounts of phosphate and potash, but

no nitrates,the amount of total fertilizeringredientin each culture
being 8o ppm. By an examinationof the table it is apparent that
the growthofeach of the creatineculturesgivenin thelast columnis
considerablylargerthan the growthof the cultureswithoutcreatine
given in the fifthcolumn. The total greenweightof the cultures
without creatine was i6. 2 grams against 23.3 grams for the cul-
tures with creatine,an increase of 44 per cent.

TABLE V
SHOWING THE EFFECT OF CREATINE ON GROWTH IN CULTURES CONTAINING NO NITRATE

PPM. OF FERTILIZER INGREDIENT IN CULTURE GREEN WEIGHT OF CULTURES

SOLUTION IN GRAMS

P2O1 NH3 K.0 Without With

creatine creatine

I . . . .. ..... 0 0 8o I.329 I*709

2 . 8
......... 0 72 I.420 1.948
3.1. - ...6
* . 0* 64
* I.558 2.I30
4 -**. . 24 0 56 I.579 2-370
5 - **. ...... 32 0 48 I.528 2.470
6... 40 . ..0 40
... . I*500 2.400
7-**. . . . ... 48 0 32 I.670 2.270
8... 56 . ..0 ...24 . I.628 2.420
9g ..... 64 0 16 i.600 2.450
TO .......... 72 0 8 I.428 2.070
II ........... 8o 0 0 0.978 I.090

In table VI are given the green weights of plants grown in

cultures with and without creatine, containing 8 ppm. of NH3
as nitrate and varying amounts of P205 and K20, the total
constituentsbeing 8oppm. of P205+NH3+K20. These figures
show that the creatine cultures given in the last column are
somewhat larger than the cultures without creatine given in
the fifthcolumn, but the differenceis not nearly so large as
in the culturescontainingno nitrate given in table V. The total
growthof the cultures without creatine was 26.4 grams against
29.4 grams for the cultures with creatine, an increase of only
i i per cent in favor of the creatine cultures. There was a
differenceof 44 per cent in favor of the creatine culturesin the
case of the solution which contained no nitrate.
i62 BOTANICAL GAZETTE [AUGUST

The growthin the cultures which contained varying amounts

of phosphateand potash and i6 ppm. of nitratewas only 3 per cent
greater with than without creatine. In solutions containing
24ppm. of nitrate the increased growthwith creatine was 6 per
cent, and in solutions containing 32ppm. nitrate the increased
growth 4 per cent. In solutions containing higher amounts of
nitratethe creatinehad no additional effect. Thus it appears that
the effectof creatinein replacingthe effectof nitratein producing
growthis very similar to that of creatinine.

TABLE VI
SHOWING THE EFFECT OF CREATINE ON GROWTH IN CULTURES CONTAINING 8 PPM. OF
NH3 AS NITRATE

PPM. OF FERTILIZER INGREDIENTIN CULTURE GREEN WEIGHT OF CULTURES

SOLUTION IN GRAMS

P2O1 NH3 KD Without With

creatine creatine

I ....... 0 8 72 2.299 2.459

2.. 8 . ..8 ..64 2.940 3.200
3... ..6 . ..8 56 2.700 3.350
4 ....... 24 8 48 2.920 3.400
5 ....... 32 8 40 3.050 3.070
6. . . . . .. .. . 40 8 32 3.I50 3.309
7 ....... 48 8 24 3.220 3.350
8 ...... 56 8 16 2.500 2.854
9g... 64 . 8
... 8 2.222 2.800
TO .t 72 .72 8 0 I.400 I.600

ect of creatineon the removal

It is also interestingto note the eff
of salts by the plants and the similaritybetween the action of
creatineand creatininein this respect. It will be rememberedthat
in the creatinineculturesthe removalof phosphate and potash was
slightlygreaterin the creatinineculturesthan the normalcultures,
but a great deal less nitrate disappeared from solution in the
creatininethan in the normal cultures.
In the creatineexperimentsthe removalof total P205, NH3, and
K20 by plants in the normal cultures was I978.3 milligrams,
against i854. 5 milligramsfor the creatine cultures. The normal
cultures removed 47 I. 0 milligramsof P205 and the creatine cul-
tures474.4 milligrams. In the case of potash the normal cultures
I9I2] SKINNER-CREATININE AND CREA TINE I63

removed769.4 milligramsof K20 against 767.4 milligramsforthe

creatinecultures. The removal of both phosphate and potash was
practically the same in the normal and creatine cultures. The
disappearance of nitratewas much less in the creatinethan in the
normal cultures. The normal culturesremoved 737.7 milligrams
against 6I 2.7 milligramsfor the creatine cultures,a differenceof
I25 milligrams.
The influenceof the creatinein regard to the removal of P205,
NH3, and K20 is very similarto that shown by creatinine,and it
again appears that this substance as well as the creatinine can
replace nitratesin its effecton plant growth.
BUREAU OF SOILS, U.S. DEPARTMENT OF AGRICULTURE
WASHINGTON, D. C.