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THE ORIGIN OF THE ERECT CELLS IN THE PHLOEM
OF THE ABIETINEAE
M. A. CHRYSLER

(WITH TWELVE FIGURES)

In the course of a comparativestudy of the phloem of conifers,
certain facts have come to light which seem to demonstratethe
originof the cells found on the marginsof medullaryrays in their
course throughthe phloem, the so-called "erect cells" of the rays.
These cells are distinguishedfromthe other cells of a ray by their
elongationin the verticaldirection,theiralmost total lack of starch,
and theirpossessionofsieve areas. Of thesethreecriteria,the third
is the finalone, while the othertwo affordpresumptiveevidence of
the nature of cells which may be found at the margin of a ray.
Such cells are characteristicof the followinggenera: Pinus, Picea,
Larix,Pseudotsuga,Cedrus,Tsuga,Abies,and also occursporadi-
callyin Juniperus
and Thrja (7).
In pittingand contentsthese cells stronglysuggesta homology
with sieve tubes, as STRASBURGER(5) has pointed out. The same
author observed that like sieve tubes these cells lose theircontents
and collapse as the season advances, while the centralcells of a ray
enlarge and become laden with starch. In his paper on ray tra-
cheids THoMPSON(7) goes so far as to say that the erect cells are
"virtually ray sieve tubes," and, fromthe fact that marginalcells
of both xylem and phloem portionsof a ray arise fromthe same
cambial cell, argues that the marginal cells in the xylem are tra-
chearyin origin. The assumptionthat erectcells are sieve tubes in
nature or originis to be put to a test in the presentpaper.
There are several possible originsfor these cells. They may,
like the ordinary(prone) cells of a ray, mergeinto the pericycle,or
theymay simplybe prone cells whichhave taken on a new function
and acquired a special shape, or theymay be cellsdistinctlyadded or
applied to a ray already existingin a simple condition, that is,
constitutedonly of prone cells.
An obvious method of attaining a solution of the question is
Botanical Gazette, vol. 56] [36
I9131 CHRYSLER-PHLOEM OF ABIETINEAE 37

suggestedby the fact that young rays altogetherlack erect cells.

This may readilybe observedin a seedlingor in the early layers of
growthof a stem or root. Attentionhas naturallybeen directedto
the stage of growthat which the erectcells make theirappearance,
and a searchhas been made among the different organsof the plant
forregionswhere a primitiveconditionwould be likely to appear.
The labors of SCOTT and of JEFFREY have shown that the
reproductiveaxis is one of the places where primitiveconditions
frequentlypersist. It has already been shown (3) that ray tra-
cheids are absent fromthe megasporangiatecone of Pinus; it may
now be added that a searchfails to reveal the presenceof erectcells
in the phloem region of the cone in Pinus, Picea, and A bies.
Accordingly,the reproductiveaxis yields no results which apply
to the presentquestion. The leaf also is not well adapted to the
present study, on account of the limited development of the
phloem. The seedling,whichhas been studiedwithsuch advantage
in other cases, has yielded results of much interestin the present
instance,especiallywhen the root is the-part used. With the roots
of seedlings have been compared young regions of roots of more
mature plants.
While Pinus has been made the basis of the present study,
material of all the genera of Abietineae except Keteleeriahas been
available. Representatives- of the hard and softpines (P. resinosa
and P. Strobus)and of species ofPicca, Larix, Tsuga, and Abies are
abundant around Orono, while material of Cedrus and Pseudolarix
has been kindly supplied by ProfessorE. C. JEFFREY of Harvard
University,and Pseudotsuga by Mr. F. D. DAvIs of Missoula,
Montana. The pines of this vicinityhave been supplementedby
P. rigida collected by Mr. L. L. WOODS of Wells, Maine, and
P. cembroides, P. edulis, and P. aristata throughthe kindness of
Dr. FORREST SHREVE of the Desert Laboratory, Tucson, Arizona.
To each of these gentlemenI wish to express my obligation. In
providingmaterial the difficulty has been to secure the necessary
stages; if the roots are too young the erect cells are absent, if too
old the cells have collapsed to such an extent as to obscure their
relationships. An age of about fourto six years has been foundto
include the significantstages in P. Strobus,but older roots must be
38 BOTANICAL GAZETTE [JULY

used in the case of nut pines. Whenever possible, the material

was preservedin a mixtureof corrosivesublimate and picric acid
acid, and
dissolvedin 30 per cent alcohol, desilicifiedin hydrofluoric
imbedded in celloidin. For renderingthe sieve areas visible,hema-
toxylin,according to EHRLICH or especiallyHAIDENHAIN, proved
useful,while a counter stain of safraninwas generallyemployed.
Russow's callus reagent was occasion-
l @|! .. ally of use.
The appearances in P. Strobus will
be firstdescribed. Fig. i representsin
radial view the phloem region of a ray
from the 6-year region of a root of a
seedlingof this age. The heightof the
erect cells is here considerablygreater
than is usually figured,for example,by
STRASBURGER (5) and also copied in his
Textbook(6), and such elongated cells
are frequent in young roots. They
suggest a close relationof these cells to
FIG. i.-Pinus Strobus, sieve tubes, as do also the sieve areas
6-year root: radial section
whichare clearly visible in this section.
throughphloem in region of a
ray, showing long erect cells The cambial regionof this and all the
withsieve pores,and a double other figuresillustratingthis paper lies
nucleusin two cases; the cam- to the right. Two of the older erect
bial regionin thisand theother g
figuresliestotheright;
X275. cells show a doublingof the nucleus,
whichhabit,as STRASBURGER points out
(5, p. 68), is characteristicof young sieve tubes. Immediatelyto
the left of these binucleate cells the crushed and empty older
marginal cells are to be seen.
Fig. 2 shows the regionborderingon a medullaryray froman
8-yearroot of P. cembroides. On the phloem side of the cambium
are to be seen a numberof prismaticcells arrangedin several series
in the radial plane. This layer of cells is intercalated between
radial rows of sieve tubes. In some of these cells sieve plates are
visible, while the protoplasm becomes more scanty as the older
regionof the phloem is approached. In other sections such cells
may be seen to lose theircytoplasmand nucleus (frequentlyafter
I9I31 CHRVSLER-PHLOEM OF ABIETINEAE 39

amitosis), and finally become completely collapsed during their

second year, afterthe mannerof sieve tubes. On the xylemside a
fewshorttracheidsare cut off,but theirwalls are thin,and theycan
be traced foronly about the width of two tracheids.
Cells in radial groupsare frequentin the phloemof youngroots
in all the species of Pinus which have been studied, and their
occurrencehas been recordedby STRASBURGER(5), who states that
they are characteristicof the young
regions of the plant, before the tan-
gential rows of phloem parenchyma I.A
cells become established. The pres-
ent investigationgoes to show that
the radial groupsof cells are not only
more abundant in the young regions,
but that they are more common in
young roots than in stems. It should
be stated however,that the cells of
the radial rows do not for the most
part correspond to phloem paren- |

chyma, as will presentlyappear. In

many cases the cambium fora period
of one or more years cuts offno cells
on the xylemside, but generallyfrom
the median region of such radial -6

groups a narrowmedullaryray is de- "'

veloped, as is clearly shown in fig.3,

whichis drawn froma 5-yearroot of FIG. 2.-P. cembroides,
8-year
P. resinosa. Cambial activity dwin- root: radialgroupofcellsprovided
dlesuntilit is confined
to themedian withsievepores; at therightthe
ofof
region
region theradialgroup,and a med- cambium
the gives
short riseto aXfew
tracheids;
~~~~~~walled
thin-
2 75.
ullaryray one cell in heightis formed.
It will be noticed that in the xylem region this ray consists of
tracheids,though cases occur where the ray cells firstformed
are tracheidsand the later ones are parenchymatous,as shown by
the pitting. Thus a tracheidalray may turn into a parenchyma-
tous one, but the reverseseems to happen seldom if ever. Atten-
tion has been called to this point by THOMPSON(7). The initial
40 BOTANICAL GAZETTE [JULy

stages in the developmentof the xylemportionof such a ray have

been observed in numerous instances, and one is representedin
fig.4, froma 7-yearroot of P. aristata. It will be seen that the
narrow ray ends on the xylem side in a somewhat pointed cell.
This method of formationof a narrow ray differsfrom the one
describedby THOMPSON in that the initialactivityseems here to be
on the phloem side, and the narrowray is wholly independentof

0 ~ ~ ~ ~ ~ ~ 0

?~~~

FIG. 3 FIG. 4

FIGS. 3, 4. Fig. 3, P. resinosa,5-yearroot: radial plate givingrise to a narrow

ray; X275; fig.4, P. aristata,7-yearroot: beginningof the productionof a narrow
ray; X275.

transitionalelementson the xylemside. Instead of givingrise to a

singlenarrowray, a radial groupmay be lined up with two or three
narrowrays, which, however,consist at firstwholly of tracheidal
cells. We have herea prolificsource of new, that is, secondaryrays
in theyoung root. I have observed the formationof parenchyma-
tous rays in the same mannerin Juniperus.
In fig. 5, fromthe same root as fig.3, is seen what frequently
takes place when one of the radial groups adjoins a medullaryray.
1913] CHRYSLER PHLOEM OF ABIETINEAE 41

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

FIGS. 6, 7. Fig. 6, P. Strobus,6-yearroot: regionbetweentwo rays,withradial

plate merginginto erect cells; X275; fig.7, 5-yearroot: two rays nearer than in
fig.6.

of these cells is shownby the sieve areas as before. On the xylem

side the cambial cell has given rise to irregularshaped tracheids
which are evidentlyof the kind describedby THOMPSON (7), and
by him regarded as transitionalformsbetween ray tracheidsand
regulartracheidsof the xylem.
It is altogetherprobable that the prismaticcells lyingbetween
two neighboringrays, as shownin figs.6-8, are of the same nature
as the cells of the radial groups,inasmuchas the contents,pitting,
I913] CHRYSLER-PHLOEM OF ABIETINEAE 43

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

o~O DOG i S~l~ali

ID
~ C 0'O

FIG. 8 FIG. 9

8, P. Strobus,5-yearroot: two rays almost in contact; fig.9,

FIG. 8, 9.-Fig.
P. aristata,7-yearroot: originof erect cells by cuttingoffa cell fromthe end of a
sieve tube; X 275.

only aftercells of a radial group become applied to its margindoes

the ray come to have a borderof "erect" cells. Such a borderis
usually added to only one edge of the ray at a time, thoughcases
occur in which bordersare added to both edges simultaneously.
A slightlydifferentmode of origin of the erect cells has been
observed in a number of cases. One of these is shown in fig. 9,
froma 7-yearrootofP. aristata. Beginningat theleftit will be seen
that the ray is not provided with a border of erect cells, but that
sieve tubes come in contact with the ray. A little fartherto the
rightan elongated cell provided with sieve pores is cut offfroma
44 BOTANICAL GAZETTE [JULY

sieve tube, and immediatelya border to the ray is produced, with

the erect cells as usual coterminous with marginal tracheids.
Similarinstanceshave been foundin otherspecies ofPinus,and the
transitionfromsieve tube to erectcell is not always as abrupt as in
the case figured. Since the cells of the radial rows are sieve tubes

\
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

of the root that the study was soon discontinued. THOMPSON'S

statementwith respectto the ray tracheidsmay be aptly quoted in
thisconnection(7, p. io8): "The rootis admittedlymoreconserva-
tive than the stem. Accordinglyin the latter the evolutionary
processesare not so well represented. There is a hurryingover of
the early stages, so that an actual seriescan rarelybe observed."
Examination of the stem of a numberof seedlingsof P. Strobus
and P. resinosashowed that (I) the rays in both the xylemand the
phloem regionswere withoutmar-
ginal cells for several years; (2)
the marginal cells as a rule make
their appearance slightlyearlierin
the phloem than in the xylem.
This feature is shown clearly in
roots as well; in fig. II, from a
S-year root of P. resinosa, is a
triangularcambial cell which has
cut offa succession of erect cells,
while as yet no cells have been
cut offon the xylem side. Fig. 7
shows the same feature,and with FIG. I P. resinosa, 7-yearroot:
it may be compared the condition the cambial cell on the lower side of
repreee ininfig. fthe raycutsoffcellson the phloem
represented 4. Addition of sidebutnotonthexylemside; X275.
the marginal cells does not begin
in all or many of the rays simultaneously,but after the plant is
two to threeyears old a fewof the rays acquire the border.
With respectto the remaininggenera of Abietineae, only a few
species have been studied with any approach to thoroughness,but
enoughhas been done to show that many of the featuresfiguredin
thispaper occurin Picea, Larix, Tsuga, and A bies. The same radial
groups of cells occur, and the phenomena seen where two rays are
verticallycontiguousmay sometimesbe seen. For instance,fig.6
mightbe almost duplicated froma young root of Tsuga canadensis.
The genus Abies presentspoints of interest,on account of the
fact that most species lack marginalcells in the xylem,thoughthey
are generallypresentin the phloem. THOMPSON pointsout that the
erect cells are "never in line with the parenchymacells of the ray,
46 BOTANICAL GAZETTE [JULY

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

sistingin the formerwhich have been lost in the latter, that is to

say, marginalcells occur in the xylemand phloem of J. communes,
while they are restrictedto the phloem of J. virginiana. If, as
recent studies indicate, the Cupressineae are derived fromlarge-
leaved coniferssuch as the Abietineae, the persistenceof marginal
cells in the phloem after they have disappeared from the xylem
must be interpretedas the retentionof an ancestral feature,which
amounts to saying that the phloem is a comparativelyconserva-
tive region.
The genusCedrusshowsin a markedway thebelated appearance
of the ray tracheidscompared with the erect cells, but a detailed
considerationof Cedrus and Pseudolarix is reserved for separate
treatmentin a futurepaper.
The choice of Pinus for detailed study has been made on the
basis of its probablyprimitivenature,in the lightof recentpaleon-
tological discoveries. If the originof the erect cells is established
forPinus, it holds forthe othergenera of Abietineae,thoughthey
may not so clearly show the formativestages. Within the genus
Pinus it appears fromthe work of BAILEY (I) that the nut and
foxtail pines are to be regarded as the most primitive,hence
P. edulis,P. cembroides, and P. aristatafromArizona were studied
with especial interest. Although difficultieswere experiencedin
judging the ages of roots, and seedlings were not available, the
material showed the same appearances as had been observedin the
eastern pines, as several of the figuresindicate. One point was
establishedwhich formsan additional argumentfor the primitive
nature of these species, namely, the marginal cells in the xylem
make theirappearance considerablylater than in the easternpines
investigated. Since marginalcells are absent fromthe reproductive
axis and from the early rings of growth in stem and root, and
particularlybecause they are not foundin the older Pityoxyla (3),
they are regardedas of comparativelyrecent introduction,and it
followsthat those species in whichtheyappear late in development
are to be consideredprimitive,unless other evidence indicates the
contrary. In respect to the-time of appearance of the marginal
cells, the nut pines and certainof the hard pines,such as P. rigid,
stand at opposite extremes,according to the limited study which
this point has received.
48 BOTANICAL GAZETTE [JULY

Doubt has been cast by BAILEY (2) on THOMPSON'S theoryas to

the originof ray tracheids (7), on the ground that the roots used
wereprobablywounded. With this criticismin mind,the material
used in the present investigationhas been carefullyexamined for
evidencesof wounds. Though the possibilityof such is not denied,
it may be said that the soundestand straightestpieces wereselected
forstudy,and no indicationsof wounds were seen in the sections.
It may be fairlyconsideredthat the relationoferectcells to sieve
tubes is established, firstbecause of the occurrencein primitive
regions of the plant of erect cells so greatly elongated as to be
eight times as long as wide, secondly because of their containing
much protoplasmbut no starch,thirdlybecause theypossess sieve
pores, fourthlybecause they eventually lose their contents and
collapse, frequentlyafterthe nucleus has divided amitotically. In
all of these respects they resemble sieve tubes. To this may be
added that theyare sometimescut offfromthe end of a sieve tube.
They are in fact sieve tubes exceptforpossessionor ratherretention
of a nucleus,forthe matterof lengthis a minorone in view of the
occurrencein the young root of much elongated erect cells, and the
greatvariationin lengthof sieve tubes in matureplants of different
species, short tubes occurringfor instance in the classic case of
Cucurbita.
In accounting for the origin of erect cells, an examination of
admittedlyprimitiveregionsof the plant seems to show that they
owe theirpresence on the edge of an originallysimple ray to the
adhesion,so to speak, ofa groupofprismaticcells lyingin the radial
plane. It has been shown that such radial groups are frequentin
young parts of the plant, especially the root, and that they occur
among radial rows of sieve tubes. But these radial plates of cells
representa passingphase; soon cambial activityin a plate dwindles
and becomes restrictedto one or a fewcells,in whichevent thereare
two possibilities: (i) one or more narrowrays may arise,consisting
on the phloem side of albuminous cells and on the xylem side of
horizontal tracheids, or (2) the cambial activity may become
localized at the edge of a ray already existing,producinga seriesof
marginalcells, the so-called erect cells. The earliermembersof a
I9I3] CHRYSLER-PHLOEM OF ABIETINEAE 49

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

are essentially short sieve tubes possessing nuclei, occurringin

groups in the radial plane. On the xylem side these may merge
into one or more narrow rays consistingof tracheids,owing to a
diminutionand localization of cambial activity.
3. When such a radial group occurs in vertical contact with a
medullaryray, cambial activity is sooner or later localized at the
edge of the ray, resultingin the productionof a borderto the ray,
such borderconsistingof sieve cells, whichare the erect cells found
in maturephloem.
4. Variationsof this mode of originof erect cells occur, such as
the cuttingof a cell fromthe end of a sieve tube when it meets the
edge of a ray.
5. In young roots and stems marginal cells may make their
appearance in the phloem earlierthan in the xylem,while in Abies
50 BOTANICAL GAZETTE [JULY

marginalcells have disappeared fromthe xylemalthoughnot from

the phloem. This and otherobservationsindicate that phloemis a
more conservativeregionthan xylem.
UNIVERSITY OF MAINE
ORONO, MAINE

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.