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THE FUNCTION OF MANGANESE IN PLANTS
WV.P. KELLEY
Historicalintroduction
Dating fromthe time of SCHEELE (i), numerousinvestigators
have noted the presenceof manganese in plants of various orders.
While small amountsonly of this elementhave been foundin most
plants, in i86o HILGARD (2) pointed out that the ash of the long-
leaf pine fromMississippi contains a relativelyhigh percentage;
and in i878 J. SCHROEDER (3) found in the ash of the Norway
spruce (Picea excels) 35.53 per cent Mn3O4,and in the ash of
the bark 4I. 23 per cent. These and otherobservations.however,
received but little attention for many years. Manganese occurs
in small amounts in practically all soils, but being an element
unessentialto growthand normal development,its absorptionby
plants was consideredto be withoutphysiologicalsignificance.
The discoveryof BERTRAND (4) in i897 of the occurrenceof
manganese in the oxidizing enzymes of plants, however,and the
subsequent findingthat small amounts of manganese salts stimu-
late the oxygen-carrying power of these catalytic agents, have
drawnattentionto thisquestionand have led to the view that after
all a physiologicalrole is probably played by this element. Since
the time of these discoveriesa large numberof experiments,with
a wide range of plants, have been made, various compounds of
manganesein water and soil culturesbeing used; and, in general,
it has been found that small amounts of manganese bring about
stimulationin growth. While small amounts oftenproduce stimu-
lation, wherevermore than a very low concentrationhas been
employed,toxicityhas resulted.
LOEW (5) and his co-workersin Japan found,furthermore, the
toxic concentrationin water cultures to be different for different
species of plants. A concentrationthat was stimulatingto rice,
for instance, proved to be toxic to barley. Likewise, SALOMONE
(6) observedin fieldexperimentsthat the applicationof such small
amounts as 4o kilogramsper hectar of manganous sulphate pro-
213] [BotanicalGazette,vol. 57
2I4 BOTANICAL GAZETTE [MARCH
observations
Experimental
The macroscopic appearance of a number of plants when
grownon manganiferoussoil is characteristic. Pineapples develop
chlorosis at an early age, fromwhich they never recover. The
effectsare firstnoticeable by a yellowingon the margins of the
leaves or the developmentof yellow spots which soon spread over
the entire leaf. The growing tips die back, the leaf margins
becoming brown. Usually such plants produce small fruitof an
abnormallypink color. Pineapple roots, instead of maintaining
theusual pointedgrowingtip,are oftenfoundto be greatlyenlarged,
sometimesto the size of a lead pencil. The roots of otherplants
also become modifiedto some extent. Those of Panicum molle
are foundto be abnormallywoody. Others are similarlyaffected.
Corn makes poor growth,the leaves turningbrown at an early
stage and the stalks taking on a deep purple color. Barley, oats,
and rice are likewisestuntedin growth. Of the Leguminosae, the
cowpea (Vigna catjang), in particular,is very sensitive to man-
ganese; the lower leaves become brown, die back fromthe tips,
and fall away. The pigeon pea (Cajanus indicus) is affectedsome-
what similarly. Onions also die back fromthe tips and produce
very small bulbs.
Some species, however,appear not to be affected. The Agave
sisalana shows no effects; sugar cane is not greatlyaffected,and
cotton and tobacco, although somewhat retarded in growth,are
not otherwiseaffected. Waltheriaamericana,the sow thistle,and
a species of Crotalaria grow as weeds on the manganiferoussoils
and are in no way hindered in their development. These illus-
trationscould be greatly enlarged, but are sufficientto indicate
that the apparent effectsof manganese in different species are far
frombeinguniform.
MICROSCOPICAL STUDIES.-From a microscopicalstudy of the
several parts of these plants, it was foundthat in those instances
where manganese exerts a toxic effect,the cell walls of the root
cortexbecome brown,and in some instancescontain minutegran-
ules of manganese dioxide. In a few instances the cells having
brown cell walls extendinto the centralcylinderof the root, and
2IS BOTANICAL GAZETTE [MARCH
TABLE I
ASH ANALYSIS OF PLANTS GROWN ON NORMAL AND MANGANIFEROUS SOILS
Phosphorica
Plants analyzed Manganese Lime Magnesia Total ash
(MnOj (CaO) (MgO) acid (PA.0s)
TABLE I-Continued
Waltheriaamericanastems:
Manganiferoussoil............. 56 I4.97 II .70 3.67 3. 58
Normal soil.................... 45 I3.70 II .30 3.46 3 73
Broom-cornleaves:
Manganiferous soil .2.24 I 2. o8 6.75 .94 ........
Normal soil................... .6o 5-44 3.52 I.38 ....
Broom-cornstalks:
Manganiferoussoil. I .36 2. I2 3. I2 I.02 ........
Normalsoil .................... 52 I.88 2.5I 3.72 ....
Tobacco stems:
Manganiferous soil. 57 9.47 4. o8 2. o8 7.02
Normal soil.................... T. I.88 3.27 7. 35 8. So
Pigeon-pealeaves:
Manganiferoussoil............. I .43 i6. II 3.25 4.04 I2. I7
Normal soil . ..........7....... 5 7.86 7.66 Io.85 7.84
Pigeon-peastems:
Manganiferous soil . . . . . . . . . . . . . 99 I5.29 2.82 4.25 5.8i
Normal soil .................... T. 3 .79 4.39 9.34 6.22
Oat straw:
Manganiferoussoil ............. .86 9. I5 5. i6 .73 I0.27
Normal soil .................... T. 3.40 3. I9 8.8I I I .73
Wheat straw:
Manganiferoussoil.22 .............2 4.5 I 4.03 2.96 8.9I
Normalsoil.................... . 2.09 2.74 5.56 i6.27
Mango leaves:
Manganiferous soil. . . . . . . . 2. i6 33. I2 2. I 2.89 9.24
Normalsoil ................ . .24 I7.13 4.90 5 .43 8.2I
practicallyeveryinstancetheabsorptionofmanganesewas increased
on the manganese soil. The analyses furthershow that thereis a
pronounced differencein the percentagesof lime, magnesia, and
phosphoricacid in the ash of plants fromthe two classes of soils.
Some of the plants were not visibly affected; others little so;
while still othersshowed a pronouncedtoxic effect;but uniformly
throughouttherewas a tendencytowardan increasedabsorptionof
lime on the one hand, and a decreased absorptionof magnesia and
phosphoricacid on the other.
Under certain conditions the ratio of lime to magnesia seems
to be of considerableimportanceto plants. In order to bringout
more clearly the relations between these two constituentsof the
ash, the relative amounts of lime and magnesia absorbed from
normal and manganiferoussoils, respectively,have been recal-
culated fromthepreviousash analysesand are presentedin table II.
An inspectionof the data in table II shows that in almost every
222 BOTANICAL GAZETTE [-MARCH
TABLE II
THE RATIO OF LIME TO MAGNESIA IN PLANTS (MAGNESIA CONSIDERED AS I)
Discussion
In this investigationit has been shown that differentplants,
whengrownon manganiferoussoils,are affecteddifferently.Some
species are stunted in growthand die back from the tips of the
leaves, which turn yellow or brown,and sometimesfall off,and a
general unhealthy appearance results. Other species appear to
be unaffected,and, so far as can be judged, vegetate normallyin
the presence of manganese. Microscopicinvestigationsshow that
in certain instances the protoplasm undergoes changes. Occa-
sionallyit draws away fromthe cell walls and the nuclei become
brown. There is a manifestchange in the protoplasmiccontents
of the roots.
The chlorophyllin a numberof plants is affected;in pineapples
it undergoesdecomposition. Simultaneouswith the destructionof
chlorophyll,starch formationceases.
The activityof the oxidizingenzymesin plants is hereinshown
to. bear no relation to the destructionof chlorophyllunder the
191I4] KELLEY-FUNCTION OF MANGANESE 2 23
LITERATURE CITED
I. SCHEELE, KARL W., Memoiresde Chymie. Dijon. I785.
2. HILGARD,E. W., Reportof Geol. and Agr.,Miss. p. 360. i86o.
3. SCHROEDER, J., Untersuchentiber Forstchem. und Pflanzenphysiol.
Tharand.Forstl.Jahrb.Supp. I :97. i878.
1914] KELLEIL-FUNCTION OF MANGANESE 227
4. BERTRAND,
G., On theoxidizingactionof manganesesalts on the chemical
composition of oxydases. Compt. Rend. 124:I355-I358. i897.
5. LOEW,O., and SAWA, S., On the actionof manganesecompoundson plants.
Coll. Agr. Imp. Univ. Tokio, Bull. 5:i6I-I72. I902.
6. SALOMONE, G., I1 manganesee lo sviluppodelle piante. Staz. Sper. Agr.
Ital. 38:IOI5-I024. I905.
7. SULLIVAN, M. X., and REID, F. R., Studies in soil catalysis. U.S.D.A.,
Bur. of Soils,Bull. 86. I9I2.
8. BERNARDINI,L., Funzione del manganese nella concimazione. Staz.
Sper. Agr.Ital. 43:2I7-240. I9O.
9. Aso, K., On the stimulatingaction of manganesechloridon rice. Coll.
Agr. Imp. Univ. Tokio, Bull. 7:449-453. I907.
10. SALOMONE, G., I1 manganesee lo sviluppodelle piante. Staz. Sper. Agr.
Ital. 40: 97-II7. I907.
ii. BRENCHLEY, W. E., The influenceof copper sulphate and manganese
sulphateupon the growthof barley. Ann. Botany 24:57I-583. I9O.
12. SCHREINER, O., and SKINNER, J. J., Organic compoundsand fertilizer
action. U.S.D.A., Bur. of Soils, Bull. 77. I9II.
13. KELLEY, W. P., The influence of manganeseon the growthof pineapples.
U.S.D.A., HiawaiiSta. Press Bull. 23. I909.
14. WILCOX, E. V., and KELLEY, W. P., The effects of manganeseon pine-
apple plants. U.S.D.A., Hawaii Sta. Bull. 28. I9I2.
I5. WooDs, A. F., The destructionof chlorophyllby oxidizingenzymes.
Centralbl. Bakt. u. Par. 5:745-754. I899.
i6. LOEW,O., and MAY, D.W., The relationof lime and magnesiato plant
growth. U.S.D.A., Bur. Plant Ind., Bull. i. I9OI; Aso, K., loc. cit., On
theinfluenceof differentratiosoflimeand magnesiauponthedevelopment
of plants. Coll. Agr. Imp. Univ. Tokio, Bull.. 4:360-370. I900-I902;
5:495-499. I903; 6:97-I02. I904-I905; LOEW,O., and Aso, K., 7:397-
407. I907.
I7. P. L., Lime-magnesia ratio as influencedby concentration.
GILE,
U.S.D.A., Porto Rico Sta. Bull. I2. I9I2.
i8. LOEB, J.,Dynamicsoflivingmatter. New York. I906.
I9. OSTERHOUT, W. J. V., On the importance ofphysiologically balancedsolu-
tions for plants. BOT. GAZ. 42:I27-I34. I906; 44:249-272. I907; On
thesimilarity in thebehaviorofsodiumand potassium. BOT. GAZ.48:98-
I04. I909.
20. , The permeability of protoplasmto ions and the theoryof antago-
nism. ScienceN.S. 35:I56-I57. I9I2; Changesin permeability
produced
by electrolytes.Science36:350-352. I9I2.