BIOLOGIA PLANTARU-M (PRAHA)

I (2) : 142--150, I959

The Effeet of Extracted H u m u s Substances on Growth

of Tomato Plants

ZDEN]~K SLADKY
Laboratory of Plant Physiology and Anatomy, Natural Science Faculty, Brno University
Received December 1, 1958

Souhrn

Jednotliv~ slo~ky humusu--alkoholick~" extrakt, huminov~ kyseliny a fulvo-

kyseliny - - postupn~ isolovan~ z kompostov~ zahradni zeminy, zbaven6 elektro-
dialysou minerAlnich iontfi, uks po p~idAni do Sachsova ~ivn~ho roztoku
rozdilnou aktivitu v pfisobeni na rdst lodyhy a ko~ene rajSat. Obdobni~ bylo po-
tvrzeno rozdiln~ ovlivn~nl pHjmu kysliku listy i ko~cny a stanoveno rfizn~ mno~-
stvl ehlorofylu v listech. Fulvokyseliny se ukAzaly aktivn~j~ii slo~kou humusu ne~
huminov~ kyseliny. Morfologick~ zm~ny ko~enov~ho syst~mu se ukg~zaly typiS-
t~j~ii pro posouzeni pdsobnosti jednotliv~ch isolovans~ch slo~ek humusu ne~ uve-
den~ zmiSny ve fysiologicks~eh charakterlstik~ch.

Summary

The different c o m p o n e n t s of h u m u s - - a l c o h o l i c extract, h u m i c acids a n d

fulvic a c i d s - - p r o g r e s s i v e l y isolated f r o m c o m p o s t g a r d e n soil a n d d e p r i v e d
o f mineral ions b y electrodialysis, were, after a d d i t i o n to Sachs's n u t r i e n t
solution, f o u n d t o have differing effects on the g r o w t h of stems a n d roots o f
t o m a t o plants. Similarly, differences in t h e effect on o x y g e n i n t a k e o f leaves
a n d roots was confirmed a n d v a r y i n g a m o u n t s o f chlorophyll were d e t e r m i n e d
in the leaves. Fulvic acids were s h o w n to be more active c o m p o n e n t s o f h u m u s
t h a n h u m i c acids. Morphological changes in the root s y s t e m were f o u n d t~) be
more t y p i c a l for a n e s t i m a t i o n o f the a c t i o n of the different c o m p o n e n t s o f
h u m u s t h a n the changes in physiological characteristics m e n t i o n e d above.

142
 E X T R A C T E D HUMUS SUBSTANCES AND GROWTH 143

Introduction

Papers dealing with the biological influence of humus on the plant (WAKS-
MAN 1936, KONONOVA 1951, 1)RXT 1954) give evidence of its far-reaching
effects on many physiological processes. Mention is made of changes in the
various processes from chemotropic irritation, stimulation, changes in plasmatic
structure, increased absorption of mineral ions, changes in the enzymatic
system, increase in growth, intensification of synthetic processes and respira-
tion, greater fertility and more rapid development. The explanation of such
a wide range of effects must be sought above all in the influence of the whole
complex of substances composing humus, with their varying biological activity
depending on the origin of the material, the method of isolation and the
sensitivity of the experimental object chosen. The data existing so far on the
biological effects of humus on plants are mainly based on the study of the
action of humus as a whole or of humic acids. Relatively little attention has
been paid to other components such as fulvic acids, hymatomelanic acids and
many others (NIKLEWSKI and WOJCIECHOWSKI 1937, KONONOVA and PAN-
KOVA 1950, KHRISTEVA 1951 and TROJANOWSKI 1954).
In order that it may be possible to make a successful approach to the
solution of important questions regarding the influence of the quality of humus
on plant growth, increased stress is placed on the necessity of studying the
action of isolated components of humus (NAJMR1957) and of a thorough
knowledge of their various modes of action (TRoJANOWSKI 1952). No study
of the complex action of humus components is, however, possible without
a detailed knowledge of anatomical-morphological changes and of the con-
sequences of changed physiological processes in the organism. It is remarkable
that very little work has been devoted to the detailed study of the morpho-
logical effects of humus substances. And yet in plants, as in all living organisms,
there exists a close connection between physiological processes and the
anatomical-morphological formation of plant organs. It was necessary to
compare the action of the different components of humus both on physiological
processes and on morphological changes and to assess whether the" effects of
these substances are similar or diverse, and whether the chemical relationship
between the humic acids and fulvic acids (MISTERSKI and LOGINOW 1957)
evinces itself both physiologically and morphologically. Therefore, experiments
were undertaken with Imun tomato plants grown in Sachs' nutrient solution
with an optimal dose of extracted humus substances (see further). Observation
of physiological changes was accompanied by the investigation of changes in
morphological structure.

Material and Methods

Fractional extraction of compost garden soil was carried out and the following humus
components were successively obtained:
1. Alcoholic extract
1. Humic acids
3. Fulvic acids
144 z. SLADK~

T h e soil was first extracted with 96~o ethyl alcohol in the cold, using a total of 75 litres to
10 kg. of soil. The alcohol was t h e n distilled under reduced pressure with a n infra l a m p (tempera-
ture 35 ~ to 40 ~ C). The residue of t h e e x t r a c t was dissolved in distilled water a t 5 0 ~ a n d
electrodialyzed b y a l l 0 V current until the i n t e n s i t y dropped below 10 mA. The yellow solu-
tion t h u s obtained was filtered t h r o u g h a blue-banded paper filter, the filtrate being t h e n used
as alcoholic extract. The soil after being extracted with alcohol was dried a t laboratory tempera-
ture, decalcified with 1~o HC1, a n d h u m i c acids were obtained b y m e a n s of preparation according
to FLAIG, SCHEFFER a n d KLAMROTH (1955). Fulvic acids were obtained b y PONO~AREV'S m e t h o d
(1947) of coagulating the acid filtrate. B o t h components, h u m i c acids a n d fulvic acids, were
electrodialyzed in the s a m e w a y as t h e alcoholic extract. The concentrations of the h u m u s
c o m p o n e n t s were determined f r o m the respective solutions b y evaporation in a v a c u u m drier
a t 80 ~ C. H u m u s substances remained in solution a n d were k e p t in a refrigerator a t 0 ~ to 5 ~ C.
D u r i n g preliminary experiments the optimal concentration of h u m u s s u b s t a n c e s in t h e
n u t r i e n t solution was established according to t h e growth of t h e t o m a t o stems. The regular
e x p e r i m e n t a l series was started on March 30, 1958 in Sachs' n u t r i e n t solution w i t h t h e addition
of t h e appropriate a m o u n t of AZ solution. The t o m a t o e s were preeultivated in n u t r i e n t solution
w i t h t h e addition of h u m u s substances in concentrations given below. A t t h e phase of two
assimilatory leaves (May 14) t h e y were transferred to litre cultivation vessels a n d tied to a silon
net. The following experimental series were set up:
1. Control plants growing only in Sachs' solution without h u m u s substances.
2. P l a n t s in Sachs' n u t r i e n t solution with the addition of h u m u s substances of optimal effect:
a) 10 rag. of alcoholic e x t r a c t to 1 litre
b) 50 rag. of h u m i c acids to 1 litre
c) 50 rag. of fulvic acids to 1 litre
There were six cultivation vessels, each containing two plants, in each series of t h e experiment.
I n all series at the s t a r t of the experiment the p H of the n u t r i e n t solution was 6.5. Differences
in t h e action of t h e h u m u s substances were recorded after 14 weeks of cultivation (July 14).
Changes in t h e height, length a n d weight of overground parts a n d roots were measured. Changes
in o x y g e n intake were followed w i t h a W a r b u r g a p p a r a t u s a n d t h e a m o u n t of chlorophyll
was determined photocolorimetrically according to GODNEV (1952).

Results

The experiments showed that the selected humus components, when added
to nutrient solution, have varying effects on the growth of tomato plants.
This is first apparent in changes in physiological processes, in more intensive
growth, and as a reflection of changed metabolic processes morphological
differences in the overground parts and root system also arise. The values
given have been statistically computed with the significance stated in table 1.
It is evident from the results that the fulvic acids are most active in their
effects on the plants. As compared with the control highly significant differences
are to be found in the height of stems, fresh weight of stems and roots and
the dry weight of stems of tomato plants grown in nutrient solution with
added fulvic acids. Only the difference in length of the root system is not
significant and in the dry weight of the roots is just significant.
A similar effect, although somewhat weaker, is to be observed with humic
acids. Only the length of the root system is greater and the difference as
compared with the control is highly significant. The alcoholic extract signifi-
cantly increased the length of stems and root system, the fresh and dry
weights of the roots and exhibited in general a less strong effect on the plants.
It was found that the alcoholic fraction increased the amount of water both
 EXTRACTED HUMUS SUBSTANCES AND GROWTH 145

T a b l e 1. D i m e n s i o n s a n d w e i g h t of I m u n t o m a t o p l a n t s g r o w i n g i n S a c h s ' n u t r i e n t s o l u t i o n
w i t h a d d i t i o n of h u m u s c o m p o n e n t s

Alcoholi c
Control Humic acids F u l v i c a c i ds
extract

H e i g h t of
s t e m in cm. 20.95 -J= 3 . 4 . 1 4 32"44 ::t= 3 9 2.33 51.52 3 9 2.23 56.76 :j:: 3 9 1.58

L e n g t h of
r o o t s in cm. 13.05 :~ 3 . 0 . 9 4 16.99 3 . l . l l 20.16 3 9 1.11 14.05 j : 3 . 1.58

Fresh
stems 6.40 =t= 3 . 1 . 5 0 8.85 ~ 3 . l ' 1 5 14.93 3 . 0.52 17.48 ~ 3 9 0"27
weight
in g.
(Gv) ro ots 1.55 =~ 3 . 0 . 3 8 3'29 3 . 0 . 4 8 3.24 3.0.51 5.39 3 . 0 . 4 6

Dry
stems 0.52 3 . 0 . 1 2 0-62 ~- 3 . 0 . 1 6 1.117 :s 3 . 0 . 1 2 1.60 -~ 3 . 0 . 1 5
weight
in g.
(Go) ro ots 0.05 ~: 3 . 0 . 0 l 0.09 :L 3 . 0 . 0 1 0.23 ~ 3 . O.07 0'24 ~ 3 9 0.07

stems 12.26 14.23 13.91 10.92

Ov/Go
roots 32.29 36.56 14.09 22.94

Difference * s i g n i f i c a n t 13 < 0.05 when P = 0.05 t = 2.06

(24)
9 * h i g h l y s i g n i f i c a n t t ) < 0.01 when P = 0.0i t = 2.80
(24)

in overground parts and roots, while humie acids caused an increase only in
the stems. Fulvic acids reduced the amount of water in the whole plant.
Changes in the oxygen intake of leaves and roots of tomato plants, as determin-
ed by the Warburg manometric technique, are recorded in table 2. It appeared
that humus substances increase oxygen intake for respiration--in the case
of overground parts fulvic acids are in the first place in this respect, followed
b y humic acids; in the case of roots the order is, alcoholic extract, fulvic
acids, humic acids. Changes in chlorophyll content of leaves of equal age are
given in table 3. They show that humus components, particularly fulvic acids
and humic acids, also increase the amount of chlorophyll.
The action of the different humus components on the overground parts and
particularly on the formation of the root system is characteristic. Figs. la to
ld show the overground parts of tomato plants from different experimental
series. As compared with the control (la) an increase in stem length and in
the sizes of individual leaves can be observed progressively from alcoholic
extract (lb), humic acids (lc) to fulvic acids (ld). Figs. 2a to 2d show the
146 z. SLADK~r

Table 2. Oxygen intake in ml. by leaves a n d roots of I m u n tomato plants per 1 rag. dry weight
per hour

Alcoholic Humic Fulvic

Control
extract acids acids

Leaves 1.757 1.934 2.183 2.283

Increase compared with control 10% 24% 30%

Roots 0.172 0.302 0.212 0.237

Increase compared with control 76% 23% 38%

Table 3. Amount of chlorophyll in mg. in 1 g. of dry weight of leaves of I m u n tomato plants

in 1 litre solvent. Average of six repetitions

Alcoholic Humic Fulvie

Control
extract acids acids

Leaves 1-213 1.581 1-983 2-054

Increase compared with control 30% 69%

differences in the formation of the root system in these experimental series

and figs. 3a to 3d illustrate differences in the branching of primary roots in
the same order. The stems of the control plants are weak at this period, with
short internodes, and are the smallest (fig. la). The upper leaves are yellowish-
green and the basal leaves fall. The root system is composed of white roots
with swollen root tips (fig. 2a). Half way along the length of the primary
roots isolated short secondary roots are to be found (fig. 3a). In the alcoholic
extract the stems are longer with short internodes (fig.lb). The leaves are
clear green and basal leaves do not fall so readily. The root system is
strongly developed (fig. 2b). The roots are long and pure white. Lateral
roots grow along the whole length of the primary root and are long and
branching (fig. 3b).
Tomato plants in humic acids form even longer stems with longer internodes
(fig. lc). The leaves are large, of a deep green colour, the basal leaves go
yellow and fall. The roots are fine, richly branched and brown (fig. 2c). The
long primary roots form fine lateral roots, mainly from the upper half; these
branch further (fig. 3c). The root tips are often destroyed.
In fulvic acids the tomato plants form the longest stems and internodes
(fig. ld). The leaves are large, deep green and those at the base also turn
yellow and fall. The root system in fulvic acids resembles that of the control
in its general aspect, b u t the number of secondary roots is greater (fig. 2d).
The primary roots are pure white, but much thicker than in the control and
with more abundant short lateral roots, mainly on the lower half (fig. 3d).
 0

a~

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tl2

0 ~

o.~

o~
 Z. SLADK~~

EXTRACTED HUMUS SUBSTANCES AND GROBrTH

Fig. 2. R o o t s y s t e m f o r m a t i o n of t o m a t o p l a n t s in S a c h s ' n u t r i e n t solution: 2a control, 2b al-

c o h o l i c extract, 2c h u m i e acids a n d 2d fulvic acids. H u m u s substances c o n c e n t r a t i o n
the same as fig. 1.
 EXTRACTED HUMUS SUBSTANCES AND GROWTH 147

Discussion

The great number of substances forming humus and their relationship to

each other considerably complicates research not only on biochemical questions
and the origin and composition of humus, but also on the question of its
influence on the plant. Contemporary chemical methods of isolation are not
adequate for the division of humus into its chemical components and therefore
the term humus substances still covers a complex of substances of the most
various chemical properties. The classification introduced by ODEN (1922),
dividing humus substances on the basis of solubility into three groups, is not
only of historical significance. I t is still in general use and formed the basis
for the isolation method of humus substances employed in the present experi-
ments.
The results of these experiments demonstrate that the presence of humus
substances in nutrient solutions causes a greater development of the root
system so t h a t the absorbent root surface is increased several times above
normal. I f we take into account the other changes in the plant which precede
the increased root development, such as changes in the enzymatic system
(SAxuBACH 1956), in metabolism (TIcH~: 1958, PR/~T, ~ATSK~ and MELICHAR
1957), changes in plasma structure (NIKL~.WSKI and WOJCIECHOWSKI 1937,
GUMII~SKI 1947), greater permeability of cell walls (LIESKE 1932, BLAGO-
WESTSCHENSKI and th~OSOROWSKAJA1934, 1935, C~AMINADE and BLANCHET
1952), we see t h a t by their complex action humus substances help the plant
to increase intake of some mineral nutrients and are beneficial in accelerating
the whole metabolism.
I t is further evident from the results that humus substances induce increased
intensity of respiration both in overground parts and roots. From the published
work of 0PXRIN and GELMAN (1952) we have evidence of a close connection
between increased respiration and increased synthesis. Other work (BANDURSKI
1957) points to an intimate connection between respiration, as a process
releasing energy, and growth, which is typified as a process of energy accumula-
tion with the co-operation of the enzymatic systems. The changes in oxygen
intake, which have been followed for this purpose, provide us with a picture
not only of respiration in the tissues, but also of the intensity of synthetic
processes, the results of which are expressed in the values for growth recorded
in table 1. Acceleration of growth, and increase in fresh weight and dry weight,
the intensity of which is typical for the action of each component of humus,
are in direct relation to the increase in consumption. There is an exception
in the case of increased oxygen consumption by tomato roots in alcoholic
extract, where this relation is not quite definite. I n agreement with BIBER
and MAGAZINER (1951) it was found t h a t fulvic acids cause more intensive
respiration than do humic acids, which undoubtedly is connected with a general
influencing of the activity of respiratory ferments (KHRISTEVA 1953).
The amount of water in the stems and roots fluctuates considerably. The
different extracted humus substances cause varying increases in dry weight
of stems and roots and point to increased synthetic activity in the plant.
This is most evident in the case of fulvic acids. The presence of humus sub-
148 z. SLADK~

stances in the nutrient solution produces an increase in and activation of the

chlorophyll apparatus, which N~KT.EWSKI and WOJCIECHOWSXI (1937) believe
to be directly influenced b y humus substances. Increased chlorophyll content
was confirmed colorimetrically as in the work of PR),T, ~ATSK~ and MELICHAR
(1957) and the amount of chlorophyll was calculated according to GODNEV
(1952). I t was found that the amount of chlorophyll increased progressively
from the series in alcoholic extract, to the humic acids and was highest with
the fulvic acids. It is evident from these experiments that the fulvic acids
are an even more effective component of humus than the humic acids and
that the influence of alcoholic extract on the processes followed is less. The
chemical relationship between humic acids and fulvic acids, established b y
physico-chemical study (MISTERSKI and LOGINOW 1957), is also clear from
their physiological action. However, morphological changes, particularly of
the root system, were found to be much more typical and reliable as indicators
of the action of the different extracted humus components. The figures show
clearly that humus substances encourage development of the root system,
the growth of which is from the start typical for the different substances. The
vitality of the whole plant is, of course, dependent on the growth and structure
of the roots.
Alcoholic extract in the nutrient solution caused increase in length of the
root system and abundant branching. The fresh weight of stems was increased
b y 38% and of roots b y 112%, mainly due to increase in water content. The
root system showed the highest oxygen consumption. This is connected not
only with increased synthetic activity in the plant, b u t also with more intense
activity of the pericambium in the formation of lateral roots.
The lengthening of roots was greatest in the humic acids. Fine, brown roots
were formed. The fresh weight of the stems was increased b y 133 ~ and of the
roots b y 109~/o, mainly due to increased water content of the stems and transfer
of assimilates to the roots, i. e. to a certain extent similarly to the process in
alcoholic extract. Fnlvic acids produced not only the greatest increase in fresh
weight of stems (170%) and roots (245%) b u t also an increase in dry weight
of 205~/o for stems and 390% for roots. The explanation is mainly to be sought
in an increase of synthetic processes and in the chlorophyll content of the leaves.
The strongly-developed root system was notable for the number of primary
roots with relatively short lateral rootlets.
I t is clear from the above that the different humus components exert
varying influences on the metabolic processes of plants and effect their physio-
logical processes with varying intensity. The action of the d fferent humus
components makes itself apparent in various forms in the general structure
of the plant, in particular of the root system.
 EXTRACTED HUMUS SUBSTANCES AND GROWTH 149

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150 z. SLADK~r

TROJANOWSKI, J.: Primenenie khromatografii dlya videleniya gumusnykh veshchestv. [The

application of chromatography in the isolation of humus substances.] - - Ann. Univ. M. Curie-
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TI~OJANOWSKL J.: Wst~pne badania had aktywnow biologiczn~ niekt6rych frakcji humusu.
[An introductory study of the activity of some humus fractious.] - - Acta Soc. Bot. Pol.
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1936.

Address: Dr. Zden~k Sladk~, Laboratory of P l a n t Physiology and Anatomy, Faculty of Natural
Sciences, Brno University, Brno, K o t l ~ s k ~ 2, Czechoslovakia.

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