Relation of Increased Water Content and Decreased Aeration to

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Agronomy & Horticulture -- Faculty Publications Agronomy and Horticulture Department
1-1930
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J. E. Weaver
University of Nebraska-Lincoln
W. J. Himmel
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Weaver, J. E. and Himmel, W. J., "Relation of Increased Water Content and Decreased Aeration to Root
Development in Hydrophytes" (1930). Agronomy & Horticulture -- Faculty Publications. 502.
https://digitalcommons.unl.edu/agronomyfacpub/502
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Plant Physiology, Vol. 5, No. 1 (Jan., 1930), pp. 69-92
Copyright 1930 American Society of Plant Biologists.
RELATION OF INCREASED WATER CONTENTA ND DECREASED
AERATION TO ROOTD EVELOPMENTI N HYDROPHYTES
J. E. Weaver and W. J. Himmel
(with eleven figures)
The dependenceo f root developmento f most plants upon aeration is
clearly shownb y water-loggingt he soil. In a few days, the usual cultivated
plants turn yellow, show wilting, and may ultimately die. If the water is
kept well aerated, plants may survive though submergedf or weeks. Ex-
clusion of oxygen from the roots of most plants interferes with the respira-
tion of the protoplasmo f their cells. This results in the death of the cells
and consequentlyt he roots fail to function as absorbersf or the plant (22).
The importanceo f aeration in plant developmenth as long been recog-
nized by plant physiologistsa nd a very large numbero f investigationsh ave
been made. In 1921, Clements (7) summarizedn early 700 such papers
in his mongraph on "Aeration and Air-Content." The long-continued
work of Cannon has added much to an understandingo f this subject and
"
his "Physiological Features of Roots (5) includes the most extensive re-
cent contributiont o this problem. The importanceo f aeration in the pro-
duction of crops and the growth of forests has been emphasizedb y the
extensive researcheso f Howard (.18), Howard and Howard (16, 17), and
Hole (14) in India. A survey of the literature shows that, with rare ex-
ceptions,t here is general agreementt hat the aeration of the roots promotes
a better growth of tops. Most of the experimentsw ere carried on in water
cultures,e .g., Hall, Brenchley and Underwood (12), Andrews and Beals
(1), and Bergman (3), and some in soil, such as those of Hunter (19),
Hole (14), Howard (15), and Knight (21). The responseo f the plant in
most cases has been measuredb y the developmento f tops and little account
has been taken, especiallyi n soil cultures, of the organs directly affectedb y
the increaseda eration. The work of Balls (2), Hunter (19), Hole (14),
and the Howards (16, 17) are amongo utstandinge xceptions. Cannon (5)
measuredt he effect of inadequate aeration on root elongation and not on
the developmento f the root system as a whole.
In devising a simple experiment for classes in plant physiology and
ecology to illustrate the effect of aeration upon root and root-haird evelop-
ment, Typha latifolia was employed (24). It was selectedb ecausei t grows
naturally under a wide range of conditions,f rom water-loggeds oil to one
that has a sufficientlyl ow water content to be well aerated. Differencesi n
root responseu nder conditions of poor and good aeration were so marked
69
70 PLANT PHYSIOLOGY
that three other characteristica nd widely distributed swamp and swamp-
margin plants were also employeda nd an extensive study of their root be-
havior begun. The use of large containers furnished an opportunity to
determinet he normal nature and extent of the undergroundp arts, a pro-
cedure which met with almost insurmountabled ifficultiesw here the plants
grow naturally in soil covered with water.
Procedure
Rhizomeso f Typha latifolia, Scirpus validus, and Spartina michauxiana
were dug in quantity on April 5, beforen ew growthh ad begun. They were
kept in a refrigeratorf or a period of 14 days before planting on April 19.
Phragmites communisw as taken from a swamp at this time and imme-
diately transplanted. Growthw as just starting.
Four large galvanized iron containers, 30 cm. square in cross section,
were used for each species. The following conditionso f soil moisture and
concomitanta erationw ere maintained: (1) saturated,n o free air; (2) alter-
nately saturated and drained, aeration poor; (3) optimum water content
for land plants, aerationg ood; and (4) dry soil, aeration good. The depth
of the containers necessary to accommodatem aximum root growth was
determinedb y preliminarye xperiments;t hose for conditions1 and 4 were
60 cm. deep, the others 90 cm. A well screened,f ertile, loam, potting soil
was used. It was mixed with one-thirdo f its volume of sand. This mix-
ture had a hygroscopic coefficiento f 10 per cent, and a water holding
capacity (Hilgard method) of about 55 per cent.
In filling the saturatedc ontainers,t he soil was thoroughlys aturateda nd
constantly stirred to permit the escape of air. After the soil had settled,
an excess of water was added to the containers o that a layer of clear water
2.5 to 5 cm. deep stood constantly above the soil. Losses by evaporation
were replacede very two or three days by adding tap water from a sprinkler.
Six liters of water were daily poured on the gravel mulch of the drained
soil. The water slowly sank through the soil mass and disappearedf rom
the surface after 10 to 30 minutes. It slowly trickled through 2.5 cm. of
coarseg ravel placed on the bottoma nd out through an opening on the side.
Here it was caught in a container and again used. This prevented loss
of nutrients or other substancesf rom the soil by leaching. Since there was
some loss by evaporation,f resh tap water was added from time to time as
needed. The third set of containers had soil of optimum water content
(25 per cent, based on dry weight) which was lightly tamped in filling.
The soil was coveredw ith a fine gravel mulch, as was the case with all the
containers except the first lot. Only enough water was added to replace
that actually lost by transpirationa nd surface evaporation. The dry con-
WEAVER AND HIMMEL: ROOT DEVELOPMENT 71
tainers were three-fourthsf illed with a dry soil which had only 12 per cent,
water content. This was but 2 per cent, in excess of the hygroscopicc oeffi-
cient. Above this was placed a 15-cm. layer of soil with a water content
of 25 per cent. Only enough water was added to the upper soil to insure
continuedg rowth.
Before planting, the rhizomesw ere carefully selected and cut in such a
manner that they were quite uniform in size, number of sprouts, etc. All
of the roots were cut close to the rhizomea nd seven rhizomesw ere planted
at depths of 3.75 to 7.5 cm. (varying with the species) in each container.
Fig. 1. Developmento f plants 17 days after transplanting the rhizomes. The sequence
from left to right is dry, saturated, moist, and drained soil.
All of the plants, except Phragrnites,r apidly developed. The growth
of the reed, althought ransplantedw hen still young, was somewhatc hecked
comparedw ith the developmento f plants in the swamp (fig. 1). For the
sake of clarity each species will be separatelyc onsidered.
Results
Typha latifolia
The developmento f the abovegroundp arts of Typha, at the end of the
35 days they were permittedt o grow, is showni n table I.
72 PLANT PHYSIOLOGY
TABLE I
Growth of Typha latifolia above ground with different moisture conditions
Maximum height, cm(cid:10)(cid:9)(cid:9)(cid:9) 111.0 106.0 86.0 66.0
Average numbero f leaves(cid:10)(cid:9)(cid:9)(cid:9) 8.4 8.1 7.6 ; 7.0
Average height of leaves, cm(cid:10)(cid:9)(cid:9)(cid:9) 46.8 | 42.3 33.0 27.5
Average width of leaves, mm(cid:10)(cid:9)(cid:9)(cid:9) 13.0 | 12.4 9.5 \ 9.4
Total dry weight of tops, grams(cid:10)(cid:9)(cid:9)(cid:9) 16.2 ! 19.9 12.2 ; 7.2
An examinationo f the table shows that in every way developmentd e-
creasedw ith a decreasei n water content. The dry weight is only an ap-
parent exception,s ince in the wet culture only 5 plants grew, but there were
7 in each of the others, except the dry soil where one plant died. Here, as
among the other three species, the plants in the moist and dry soils had a
darker green color throughoutt heir period of growth, due probably,t o the
more favorable conditions for nitrification and nitrogen fixation. In the
saturated and drained containerst he roots were readily freed from adher-
ing soil particles and their dry weight was obtained. The dry weight of
tops and roots was in the ratio of 5.8 to 1 in the saturateds oil and 5.6 to 1
in the drained.
Saturated culture. - The root systems were examined on May 23-25
when the plants were five weeks old. Thosei n the saturateds oil were very
shallow and consisted of two distinct parts, namely, soil roots and water
roots. All originatedf rom a basal node of the stem about 6 cm. below the
soil surface and none from the old rhizome. The rhizomess hriveled upon
the removalo f accumulatedf ood and in several cases they were decompos-
ing, although the soil was not sour. The soil roots, 64 in number on one
of the largest plants, pursued courses at all angles between the horizontal
and the perpendiculari n such a manner as to more or less thoroughly
ramify a hemisphereo f soil with a radius of 15 cm. Eleven of the long-
est extended to 28 cm. depth.
These shining white roots were 1.5 to 2 mm. in diametera nd very turgid
and brittle, especially the distal 10 cm. Branchesw ere entirely absent on
most of the roots or occurredo nly very rarely. On others, slightly brown-
ish in color and probablyo lder, thread-likel aterals arose at the rate of 1 to
8 per cm. (rarely 14 on the best branchedp arts). These were exceedingly
variable in their distribution, frequently long portions of the roots being
bare. Laterals were 1 to 5 cm. (maximum1 0 cm.) long, and entirely un-
branched but much more abundantly furnished with root hairs than the
main roots wheret hey were relatively sparse.
WEAVER AND HIMMEL : ROOT DEVELOPMENT 73
Fig. 2. Typha latifolia grown in soil covered with standing water, showing the sharp
differentiation between soil and water roots. Photograph taken with
root system under water in its natural position.
Approximatelyh alf of the root system (66 roots by actual count) con-
sisted of water roots. These arose at a higher position on the node than
most of the soil roots. While some ran obliquely upward, often at a 45°
angle, many grew vertically and then turned at right angles after leaving
the soil (fig. 2). Other roots ran horizontallya bout 8 cm. and then grew
abruptly upward. The portions in the soil were like those described,b ut
upon entering the water they greatly decreasedi n diameter and branched
profusely, the distal part becoming extremely attenuated (figs. 3 and 4).
They began to appearo nly 5 days after the rhizomesw ere planted and their
growth was rapid, some reachinga length of 10 cm. in 3 days. By the ex-
piration of 10-14 days all were greatly branchedw ith laterals a centimeter
or less in length. New roots continuallya ppeareda nd branchingi ncreased
throughout the period of growth. The older ones reached a maximum
length of 30 cm. but many of them were only about one-halfa s long. When
mature, they were branchedt o near the tips but there were no laterals of
the second order. The very fine primary laterals, which occurreda t the
rate of 20 to 55 per cm., varied in length from 1.5 to 2.5 cm. and were en-
74 PLANT PHYSIOLOGY
Fig. 3. Water roots of Typha about two weeks after transplanting the rhizomes.
tirely free from root hairs. Thesef ine water roots and their very abundant
branchese normouslyi ncreasedt he absorbings urface in the aerated portion
of the culture. Aside from frequent renewal of the water, an abundance
of Ulothrix and Mougeotiai ncreasedt he oxygen content. The former alga
was so abundantlya ttached to the roots that it gave them a greenish hue.
Drained culture. - The root system in the drained soil was much
deeper seated althought he roots were most abundanta nd most branchedi n
the surface 9 cm., which was the best aerated. Here they formed a dense
Fig. 4. Detail of water root of Typha. Note the attenuated tip and the profuse simple
branching.
WEAVER AND HIMMEL : ROOT DEVELOPMENT 75
network. Roots were numeroust o a depth of 30 cm. and some attained
depths of 40 to 47 cm. (fig. 5). The general distributiono f the roots was
such as to occupy a more or less hemisphericalm ass of soil. They were
rather clearly differentiatedi nto three types as regards position, thickness,
and degree of branching.
Pig. 5. Typha grown in drained soil. The finer, horizontal roots are near the soil
surface.
One lot consistedo f coarse,u nbranchedo r practically unbranchedr oots
which, as a group, penetratedd eepest. Their diametersw ere 1.5 to 2 mm.
throughout. A second lot was composedo f roots with smaller diameters
(1-1.5 mm.), some of which penetrated as deeply, or at least nearly as
deeply, as the former. These branchedp rofusely, often forming networks
of laterals at depths of 30-36 cm. where branches1 0 cm. long were found.
The branchingh abit was variable; on some, only the proximal portion was
76 PLANT PHYSIOLOGY
branched,o n others,t he distal part. On still otherst he presenceo f branches
and especiallyt heir density varied from place to place. The rate of branch-
ing was 6 to 25 laterals per cm. The branchesr anged in length from less
than 1 cm. to a maximumo f 10 cm., althoughm ost of them did not exceed
3 cm. Nearly all of the primaryl aterals were simple; a few were branched
to the first order, the branchess eldom exceeding 3 or 4 per lateral, and a
centimeteri n length.
Fig. 6. Detail of one of the surface roots of Typha grown in drained soil.
A third type of root arose from the highest portion of the node at the
base of the stalks and thus above the other roots. They thus originateda t
a depth of about 7 cm. and no portion of the root extended deeper than the
point of origin. In fact, a few at first extendedv ertically upward but this
tendencyw as not nearly so markeda s in the case of water roots in the pre-
vious culture. Brancheso f these fine roots (which were 0.5 to 1 mm. thick)
often occurred throughout,b ut in general they were most abundant and
WEAVER AND HIM MEL : ROOT DEVELOPMENT 77
longest on the distal one-thirdt o one-half. Laterals were 7 to 10 per cm.
on the more thinly branched parts and 12 to 20 on those most thickly
branched. Theset hread-liker oots were shining white, and the longest were
more abundantlyf urnished with laterals than were any in the deeper soil.
Secondaryb ranchesw ere few. The main root and all its branchesw ere well
supplied with root hairs. These roots differedf rom the water type by .hav-
ing longer, coarser,a nd more crinkly laterals with some sublateralsa nd an
abundanceo f root hairs (fig. 6). The last was in striking contrast to the
water roots, which had none.
One of the larger plants had 56 of the deep, unbranchedt ype, 33 of the
deeply penetrating branched ones, and 43 of the fine, surface type which
were most favorablys ituated as regards aeration.
Moist culture. - The root system in the moist, well aerated soil showed
no differentiationi nto parts ; all of the roots were 1 to 1.5 mm. in diameter
and about equally branched. There were no roots in the surface 7 cm. of
soil. Long horizontalo nes ran outward, sometimest o distances of 30 cm.,
but never turned downward. Othersr an outwarda nd downwardt o depths
Fig. 7. Relative branchingo f roots of Typha; A, in saturated soil; B, in wet but drained
soil; and C, in moist, well-aerated soil.
of 40 to 53 cm. All, except the shortest,w ere regularlyb ranchede xcept the
rapidly growing,s hining white distal 8 cm., upon which laterals had not had
time to develop. Branches occurred at the rate of about 8 per cm. and
varied in length from 1 to 2.5 cm. Secondaryb ranchesw ere only 1 to 3
mm. long and occurreds parsely and only on the longest primary branches
(fig. 7).
Dry culture. - The root system was very much like that in the moist
soil but more abbreviated. Its extent was determinedb y the depth of the
moist soil which varied from 16 cm. on one side of the containert o 35 on the
other. None occurredi n the surface 9 cm. (fig. 8). The dwarfed plants
had fewer roots (27 and 51 respectivelyo n two of the largest plants) than
in moist soil but they were much more densely branched,e .g. 10-35 laterals