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El Origen de El Erect Cells
El Origen de El Erect Cells
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THE ORIGIN OF THE ERECT CELLS IN THE PHLOEM
OF THE ABIETINEAE
M. A. CHRYSLER
0 ~ ~ ~ ~ ~ ~ 0
?~~~
FIG. 3 FIG. 4
The cells in question apply themselvesto the side of the ray, and as
cambial activity in the radial group slackens, this activity is
restrictedto the borderof the ray, thus changingthe ray froma
simpleone consistingonlyofparenchymatouscells to a ray provided
with marginal cells. With fig. 5 may now be compared fig. I,
which shows the same mode of originfor the marginalcells of the
phloem regionof the ray in P. Strobus. Observationshave been
made on representativesof the hard and soft pines, includingthe
nut and foxtailpines, and appearances similar to fig.5 have been
found in all, when roots of appro-
priate age were examined.
Regionswheretwo rays lie near
one another in the vertical plane _
present appearances which may
readilybe explained in the lightof
what has been said of the radial
plates. Fig. 6 shows such a region
in a 6-year root of P. Strobus. In
the upper part of the figureis a
ray which on one margin is desti- 4
tute of erectcells, and on the other
(lower) marginis provided with a A
fringe of much elongated cells
which soon merge into a group of FIG. 5. P. resinosa, 5-yearroot:
prismatic cells forming a radial a radial plate is in contact with a
ray,producinga row of
of tissue
plateof
plate tissue by means of
by means which medullary
of which marginalcells; X275.
the ray shown is connected with
another which lies below the area covered by the figure. These
cells of the "radial plate" show precisely the same histological
featuresas thosepreviouslydescribed,such as the sieve areas which
are hereshown. At the extremeleftof the figureis a rowofphloem
parenchymacells, which are easily distinguishedfromthe cells of
the radial plate by the starchy contents and swollen shape of
the former.
Furthertransitionstages are shownin fig.7, froma 5-yearroot
of P. Strobus,whichrepresentscases wheretwo rays are closer than
those in fig. 6. Almost from the time of their formationfrom
42 BOTANICAL GAZETTE [JULY
cambial cells the erect cells of the two rays come into contact, and
soon merge into the conditionwhere a single cell spans the space
between the rays. As before,such cells lose their contents and
become greatlycompressedin the radial direction.
Fig. 8, fromthe same root as fig.7, representsa case where two
rays are practicallyin contact. A cambial cell is seen to give rise
on the phloem side to pairs of cells which appear to have earlier
been undivided,as seen in the extremeleftof the figure; the nature
FIG. 6 FIG. 7
and fate of the cells are the same in both cases. Where two rays
are in verticalproximity,it is to be expected that the intervening
space will be occupied by one or two cells, but here,as in the case
of the radial plates of cells, thereis seen a tendencyforthe cambial
activityto be localized in the regionof a ray.
Several of the figures illustrate a point which must be
emphasized,namely,a ray uniformlybegins its course in the outer
phloem as a simple structureconsistingexclusively of '"prone"
cells which at certain times of year contain abundant starch,and
ID
~ C 0'O
FIG. 8 FIG. 9
\
in all respectssave lengthand the nucleus,the occasional formation
of an erectcell by cuttingoffa segmentfroma sieve tube does not
seem surprising. In fact it does not seem necessaryto assume an
1 I absolutely
uniform
modeofori-
gin for the erect cells, in view
ofthe factthat a certaincambial
cell may be giving rise to ray
tracheids,but for some reason
suddenly cut offa parenchyma
cell instead. Similarlya row of
ray cells of the phloem may at
firstbe albuminous, but later-
formedmembersof the rowmay
contain starch. We are inter-
ested here,however,in the ques-
tionas to the evolutionaryorigin
of the erect cells, ratherthan in
FIG. io.-P. Strobus,4-year root: occasional modes of origin.
sieve tubes have cut offcells fromtheir An interestingmode of for-
in production
ends,resulting of a ray;-
X275. 2 ~~~mation
of a ray, showingthe
X275.
tendency of sieve tubes to cut
offa cell fromthe end, has been observeda fewtimes. An example
of this mode, froma 4-year root of P. Strobus,is representedin
fig.io. In this case the habit of cuttingoffa cell fromthe end of
a sieve tube has become so well established that a narrowray has
originated,as seen in the cambial regionat the right. This process
is entirelysimilar to the mode of originof erect cells describedin
the precedingparagraph.
Althoughthe most significantresultshave been obtained froma
study of the young root, the stem of Pinus has also been examined
to some extent,but the mode of originof the erectcells in this case
was so indefinitecomparedwithwhat may be seen in similarregions
1913] CHRYSLER-PHLOEM OF ABIETINEAE 45
but always above or below them. Often they are in line with two
or three degeneratingcells on the wood side" (7, p. II 2). The
conclusionseems justifiedthat Abies representsa genus which has
descended fromancestorswhichpossessed ray tracheids,and is less
primitivethan Pinus. In spite of this,it may sometimesbe made
out in young roots of Abies balsamea that the erect cells have the
same mode of origin as has been described for Pinus. Fig. I2
illustratesthispoint,as well as the fact that the triangularcambial
cell gives rise to erect cells but not to ray tracheids. In many
instances the shadowyremainsof marginal
tracheidsappear in these root sections.
i.l o
i With thispersistenceofmarginalcells in
the phloem may be compared theirearlier
appearance in the phloem than in the
xylem,as seen in seedlingsof Pinus. Just
C)rwhy a cambial cell should be more apt to
cut offsegmentson the phloem side than
on the xylemside is hard to say; since the
supply of food is on the phloem side, un-
> equal nutritionmay be the cause at work.
But it is possible that we must considerthe
FIG. I2.-Abies balsamea, phloem to be a more conservative region
8-yearroot:theerectcells
have the same mode of than the xylem,in which case the observa-
origin as in Pinus; the tion has interest from the phylogenetic
cambial cell at margin of standpoint. The latter view of the case
the ray cuts offerect cells
butnoray tracheids; 275.is supported by my observations on Jf-
niperus, in which genus PENHALLOW (4)
found ray tracheidsoccurringsporadically. If a radial section of
J. communisis cut so as to includephloemas well as xylem,it is seen
that whereray tracheidsoccur near the cambium they are cotermi-
nous with erect cells. But in the root of J. virginiana,in which
species ray tracheidshave not been reported,a numberof instances
have been foundwherethe phloemportionof a ray is providedwith
a border of well marked erect cells, ending at the cambium in a
blunt cell, remindingone of the appearances in Abies balsamea.
Judgingby the leaves, J. communisis a moreprimitivespeciesthan
J. virginiana,hence we are prepared to findancestral featuresper-
19131 CHRYSLER-PHLOEM OF ABIETINEAE 47
seriesof these cells are more elongated than the ones formedlater,
but all possess the same histologicalfeaturesand undergothe same
fate. Similar in formand contents are the cells which span the
space between two verticallycontiguousrays, and which by fission
and shorteninggive rise to a row of erect cells on each of the
two rays.
Since the cells of the radial rows occur principallyin the young
plant, and have the same contentsas sieve tubes,withthe exception
of the nucleus, it is a temptingtheoryto consider them the fore-
runnersof sieve tubes, the primarysieve cells whichundergoevolu-
tionin two directions: (i) lengtheningout and losingthe nucleusso
as to producesieve tubes,and (2) shorteningas theybecome applied
to medullaryrays and become convertedinto erect cells or as they
give rise to rays independently. But such a theorymust stand on
the evidence of a comparativestudy of phloem of vascular plants,
and the evidence is not yet at hand.
Summary
ir. The "erect cells'" occurringon the marginsof medullaryrays
in the phloemofmostgeneraofAbietineae do not existin the young
ray, which consistsonly of ordinaryparenchyma.
2. In youngrootsof Pinus the phloem shows certaincells which
LITERATURE CITED
I. BAILEY, I. W., Anatomicalcharactersin the evolutionof Pinus. Amer.
Nat. 44: 284-293. pI. 2. I9I0.
2. , A Cretaceous Pityoxylonwith marginal tracheids. Annals of
Botany 25:3I5-325. PI. 26. I9II.
3. JEFFREY, E. C., and CHRYSLER, M. A., On Cretaceous Pityoxyla. BOT.
GAZ. 42: I-I5. pIs. I, 2. I906.
4. PENHALLOW, D. P., A manual of the North American Gymnosperms.
Boston. I907.
5. STRASBURGER,
E., tUberden Bau und die Verrichtungender Leitungsbahnen.
Jena. i8gi.
6. ---, Lehrbuch der Botanik. Neunte Auflage. Jena. I908.
7. THOMPSON,
W. P., The originofray tracheidsin the Coniferae. BOT. GAZ.
50:ioi-ii6. I9IO.