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 THE ROLE OF OXYGEN IN GERMINATION
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY i8i
CHARLES A. SHULL

A study of the respirationof Xanthium seeds was undertaken

some time ago with the purpose of determiningwhethertherewas
any change in the permeabilityof the seed coats to oxygenduring
the period followingthe normal ripening of the seeds. Some
evidence was noted previously that there was either a change in
permeabilityof the seed coat, or a change in oxygen need of the
embryoduringthe early winter,and it was believed that a careful
measurementof the oxygenused by the seeds with coats on and
offat successive intervals during the year would show which of
these changesoccurred,and at what period of the ripeningprocess.
Circumstanceshave prevented the carryingout of this series of
tests; but the preliminaryresultsare of sufficientinterestin con-
nectionwith the role of oxygenin germinationbehavior to warrant
placing the data on record. The measurementswere made with
a respirometer ofexcellenttypedesignedby Dr. WILLIAMCROCKER,
to whom I am furtherindebted for suggestions regarding the
problem. The respirometerwas kept in a Freas thermostatat
25. 25? C., and the volumes of oxygenused are reduced to standard
conditions. Seeds of X. glabratumin dry storage fornine months
were used.
First it was necessary to know what part of the oxygen was
used by the coats under ordinaryatmosphericgerminativecon-
ditions. Two lower seeds were placed in one chamber of the
respirometer,and the coats of two lowers in the other chamber.
In 22.5 hours the two seeds used 0.475 cc. of oxygen,while the
two coats used o. o98 cc. From the resultsof BECQUEREL'Swork2
I had suggestedthat the coats were probablyresponsiblefora part
I SHULL, CHAS. A., The oxygen minimumand the germinationof Xanthium

seeds. BOT. GAZ. 52:453-477. 1911.

2 BECQUEREL, PAUL, Recherchessur la vie latente des graines. Ann. Sci. Nat.
Bot. IX. 5:193-320. 1907.
BotanicalGazette,vol. 57] [64
I914] SHULL-GERMINATION 65

of the respirationof intact seeds. That part is now shown to be

considerable,amountingin thisinstance to 20 per cent of the total.
The respirationof lower and upper seeds with coats intact
under atmosphericconditionsof germinationwas compared, with
the result that two lowers used 0.687 cc. and the two uppers
0. 509 cc. of oxygen in 42.3 hours, a ratio of lowers to uppers of
I.35: I. It should be said that upper seeds always weigh less
on the average than the lowers; and in using equal numbersof
seeds the weight of respiringsubstance is somewhat less in the
uppers.
The respirationof the lowers with coats on and coats offin
ordinaryatmosphereis especiallyinteresting(see table I).
TABLE I

OXYGEN USED
TIME RATIO
Coats off Coats on

I0 hours 0.. 500 cc. 0.36I5 CC. I *38: I

I-I*5 0.478 0. I025 4. 65: I
I5-I7 . 25 * 0.3I6 " . 040 7.9 :I

Total. ......... I.294 0- 504 2.57 I

TABLE II

OXYGEN USED
TIME RATIO
Coats off Coats on

5 hours
....... ... o6 cc. 0 .075 CC. 2. I4: I
it . . ........ o. 6i8 " 5 58: I
5-I 5 0 0.II07 *
Total, I 7. 25 hours... o. 9825 0 I947 5.046: I

A similar test with uppers is shown in table II. The coats were
not placed in the chamberswith the embryoswherecoats were off,
so that the ratiosin the tables are too low as regardsactual embryo
respiration. The rapid increaseofuse of oxygenby naked embryos
as germinationcommencesis well illustrated.
Finally, the oxygen used by uppers and lowers with coats on,
in an atmosphere96 per centoxygen,was compared,withsignificant
results. Two lower seeds used I .007 cc. in I 2. 5 hours, while the
66 BOTANICAL GAZETTE [JANUARY

uppers used but 0.4406 cc., a ratio of 2.28:i. A repetitionof

this experimentresultedin the lowersusing I . 257 cc. in 22 hours,
theuppers0. 772 cc., a ratioof I. 63: i. Invisibledefectsin coats
might cause some variation in these ratios, but they are believed
to approximateaverage results.
It is interestingto note that two lowers in atmosphereused
o.687 cc. of oxygen in 42 .3 hours, while the same number used
I.007 CC. in I2.5 hoursin 96 per cent oxygen; and that uppers
which used o . 509 cc. in 42 .3 hours used o . 4406 cc. in I2.5 hours
in 96 per cent oxygen. The increased oxygen pressure causes a
large increasein the oxygenintake of both seeds with coats intact,
but exerts the greater influence on the lowers. The relation
between oxygen influenceand respirationseems to be close. At
least we may say that the conditionsof the oxygensupply which
lead to increaseduse of oxygenare just the conditionswhichbring
about germination. The possibilitythat oxygenexertsits stimu-
lative effecton germinationby increasing respiration,thereby
yielding more energy, is stronglysuggested, without, however,
precludingthepossibilitythatothereffectscorrelatedwithincreased
respirationmightdetermineits influencein germination.
BECKER3 recentlytested the influenceof oxygen on the germi-
nation of seeds of several plants. The fruits of Dimorphotheca
pluvialis were found to germinatemore readily in 02 than in air,
the ray seeds especially showing the favorable influence. Short
exposuresto oxygen (I 5 hours) had no such effect,but if the time
of exposure were lengthenedto 30 hours, this exposure favored
furthergerminationunder atmosphericconditions. The ray seeds
again showed the effectmore stronglythan the disk seeds. When
the fruit and seed coats were removed, io hours' exposure to
oxygen affectedgerminationfavorably,but I3 hours' lengthened
the germinationtime. The seeds of Calendulaeriocarpawith coats
intact were greatly favored by oxygen, while the yellow-brown
verticalfruitsofA triplexhortensis and A. nitensshowedan injurious
effectfromincreasedoxygenpressures. The relationof oxygento
germinationin these cases seems to be irregularand inconstant.
3BECKER, H., Uber die Keimung verschiedenartiger Friichteund Samen bei
derselbenSpecies. Beih. Bot. Centralbl.29 :21-143. 1912.
I9I4] SHULL-GJ?RMINATION 67

BECKER draws a generalconclusionfromhis resultsthat oxygen

acts as a stimulus, and takes particular exception to the idea
advanced by CROCKER4 that the oxygen increases the respiration
and in this way initiates germination. In reviewingBECKER'S
paper,5I stated that there was no doubt that in Xanthium the
oxygen was actually used in germination,and suggested that
increased respirationmight be identical with BECKER'S stimulus.
While it is entirelypossible that the oxygen influenceis exerted
throughsome other process correlatedwith increased respiration,
the data presentedhere give the ground upon which that sugges-
tion was based. Unfortunately,BECKER'S work gives us no data
as to the respirationbehavior of the seeds on which he worked,
so that no comparisonwith the behaviorof Xanthiumseeds can be
made at present.
LEHMANN,6in discussingthe possibilitythat 02 mightact as a
catalyst, accepts BECKER'S idea that oxygen acts as a chemical
stimulus, not merely by increasing respiration. Of course, the
word " stimulus" is vague and indefinite. But it should not
implyan additional absorptionof oxygen,forthis could not occur
withoutinvolvingoxidationof some kind, whichwould be respira-
tion. Even if oxygenis conceived to be a catalyser,that concep-
tion does not involve increased use of 02, for catalysts are not
used in the processesthey carryon.
The biological role of oxygen is so complex that we may not
say its effectis always due to increasingrespirationor oxidation.
The role, indeed, may be different in differentseeds and plants.
For instance, ARPAD PAAL7 claims that reductionof oxygenpres-
sure even to a. 75 normal lengthens geo-presentationand geo-
reaction time to a marked degree. This work of PAAL'S still
awaits confirmation. And although the earlier work of STICH,
JOHANNSEN,and others indicates that this amount of reduction
4 CROCKER,WM., The r6le of seed coats in delayed germination. BOT. GAZ.
42:265-29I. i906.
5 BOT. GAZ. 54:433. I912.

E., und OTTENWALDER, A., Uber katalytischeWirkungdes Lichtes

6 LEHMANN,

bei der Keimunglichtempfindlicher Samen. Zeitschr.Bot. 5:337-364. I9I3.

7 PAAL, ARPAD, Analysedes geotropischen
ReizvorgangsmittelsLuftverdunnung.
Jahrb.Wiss. Bot. 50:I-20. I9II.
68 BOTANICAL GAZETTE [JANUARY

should not affectthe rate and nature of respiration,yet it would

be very desirable to repeat PAAL'S work, studying the rate of
respiration along with presentation time and reaction time to
discover whether there is a parallel effectwith that on the res-
piration rate and germinationpower of seeds.
ZALESKI8 has noted the influenceof oxygen on the rate of
protein synthesis. IVANOFF9 has shown that oxygen is necessary
for the transformationof zymogen into zymase, and, as is well
known,thereare a numberof oxygencarriersand oxygenabsorbers
in the livingcell. Xanthophylland otherpigmentsabsorb oxygen,
lecithinplays a similarrole, and PALLADINI? has now shown that
his respiratorychromogenstake up oxygen readily. All of these
factsgo to show how complexthe oxygenrole may be, and suggest
some of the possibilitiesof even briefexposureof seeds to oxygen.
On the otherhand, however,it would be strangeif the oxygen
effectin some cases were not due simply to its influenceupon
respiration. The influenceof the amount of oxygen present on
aerobic and anaerobic respiration,which differso markedlyin the
amount of energyreleased,is well known. Anaerobic may change
over to aerobic on access of oxygen,with a consequent rapid rise
in energyreleasal leading to germination.
With these Xanthium seeds it has been shown that when the
oxygensupply is increased,it in some way bringsabout an imme-
diate and rapid increase in the rate of oxygenabsorption. At the
same time,the increasedoxygensupplybringsabout an immediate
germinationof the seeds. The two effects,increased absorption
and germination,are closely correlated as regards their relation
to the oxygen supply. This shows conclusively,I believe, that
the assumptionmade by BECKER and LEHMANN, which led them
to reject the idea that the influencewas exertedthroughincreased
respiration,is not correct,so far as Xanthiumis concerned. Nor
does theirworkthrowany lighton thisparticularpoint, since they
8ZALESKI, W., Zur kenntnisder Stoffwechselprocessein reifendenSamen. Beih.
Bot. Centralbl.27:63-82. I9II.
9IVANOFF, L., Uber die sogenannte Atmung der zerriebenenSamen. Ber.
Deutsch. Bot. Gesells. 29:563-570. I9II.
IOPALLADIN, W., and TOLSTAJA, Z., Uber die Sauerstoffabsorption durch die
Atmungschromogene der Pflanzen. Biochem.Zeitschr.51:38I-397. 1913.
19I4] SHULL-GERMINA TION 69

did not measure the respirationof the seeds on whichtheyworked.

However, owing to the complexityof the oxygen role in physio-
logical processes,it is very difficultto say just which functionor
functionsare affected. It seems certainthat the oxygenacts as a
limitingfactoron some function,whetherby limitingthe process
of respirationor energy releasal, by limitingenzyme formation
or the action of oxygencarriers,or in otherstill less definiteways.
The exact methodby which absence of oxygendelays germination
can be determinedonly by furtherinvestigation. In the mean-
timetheoriesmay well await the factswhichwillmake philosophical
discussionof this question unnecessary.
UNIVERSITY OF KANSAS
LAWRENCE, KAN.