Panzer Divisions 194445 Original Pier Paolo

Battistelli download

https://ebookbell.com/product/panzer-divisions-194445-original-
pier-paolo-battistelli-4449564

Explore and download more ebooks at ebookbell.com

Here are some recommended products that we believe you will be
interested in. You can click the link to download.

Panzer Divisions 194445 Pier Paolo Battistelli

https://ebookbell.com/product/panzer-divisions-194445-pier-paolo-
battistelli-6757098

Panzer Divisions 194445 Pier Paolo Battistelli

https://ebookbell.com/product/panzer-divisions-194445-pier-paolo-
battistelli-11924200

Hitler Youth And The March Of The Ss Panzerdivision 194445 Warrior Im

Baxter

https://ebookbell.com/product/hitler-youth-and-the-march-of-the-ss-
panzerdivision-194445-warrior-im-baxter-2166234

Panzer Gunner From My Native Canada To The German Ostfront And Back In
Action With 25th Panzer Regiment 7th Panzer Division 194445 Bruno
Friesen

https://ebookbell.com/product/panzer-gunner-from-my-native-canada-to-
the-german-ostfront-and-back-in-action-with-25th-panzer-regiment-7th-
panzer-division-194445-bruno-friesen-52705958
Panzer Lehr Division 194445 Fred Steinhardt

https://ebookbell.com/product/panzer-lehr-division-194445-fred-
steinhardt-37745938

Panzers On The Eastern Front General Erhard Raus And His Panzer
Divisions In Russia 19411945 Peter Tsouras

https://ebookbell.com/product/panzers-on-the-eastern-front-general-
erhard-raus-and-his-panzer-divisions-in-russia-19411945-peter-
tsouras-33693518

Panzers On The Eastern Front General Erhard Raus And His Panzer
Divisions In Russia 19411945 Tsouras

https://ebookbell.com/product/panzers-on-the-eastern-front-general-
erhard-raus-and-his-panzer-divisions-in-
russia-19411945-tsouras-6636076

Panzers On The Eastern Front General Erhard Raus And His Panzer
Divisions In Russia 19411945 Pdfdrivecom Mobi Peter Tsouras

https://ebookbell.com/product/panzers-on-the-eastern-front-general-
erhard-raus-and-his-panzer-divisions-in-russia-19411945-pdfdrivecom-
mobi-peter-tsouras-33693606

Panzer Divisions At War 19391945 Images Of War 1st Edition Ian Baxter

https://ebookbell.com/product/panzer-divisions-at-war-19391945-images-
of-war-1st-edition-ian-baxter-46867360
Battle Orders
OSPREY
PUBLISHING

Panzer Divisions
1944–45

16

le
Se
KL
Se
II 71 129 29
16
I 71
KRÜGER ULICH
KL
KRÜGER
III 71

50
KRÜGER
II 143
Ca lor e 100 I 15
ULICH
50

III 143

Altavilla
I 142 II 15
151
ULICH

III 142
751
1

Pier Paolo Battistelli • Consultant editor Dr Duncan Anderson

© Osprey Publishing • www.ospreypublishing.com
Battle Orders • 38

Panzer Divisions
1944–45

Pier Paolo Battistelli

Consultant Editor Dr Duncan Anderson • Series editors Marcus Cowper and Nikolai Bogdanovic

© Osprey Publishing • www.ospreypublishing.com

 Contents

Introduction 4

Combat mission 5

Doctrine and training 6

Unit organization 8
From 1943 to 1944 • Divisional Headquarters • The Panzer Regiment • Panzergrenadier units
Panzeraufklärungs Abteilung • Panzerjäger Abteilung • Artillery • Panzer Pionier Bataillon
Communications and replacements • Divisional supply and services • The 1945 Panzer Division

Tactics 46
Salerno, 13–14 September 1943 (16. Panzer Division) • Gorodok, 21–27 November 1943 (20. Panzer Division)
Cherkassy, 16–17 February 1944 (1. Panzer Division) • Targul–Frumos, 2 May 1944 (24. Panzer Division)
St. Lo, 11 July 1944 (Panzer Lehr Division) • Caen, 18 July 1944 (21. Panzer Division)
Arracourt, 25–29 September 1944 (11. Panzer Division) • Nyiregyhaza, 26 October 1944 (1. and 23. Panzer Divisions)
Hürtgen Forest, 2–9 November 1944 (116. Panzer Division)
Celles–Rochefort, 23–26 December 1944 (Panzer Lehr, 2. Panzer Division)
Wissembourg, 8–9 January 1945 (21. Panzer Division) • Stuhlweissenburg, 20–21 January 1945 (1. Panzer Division)

Weapons and equipment 70

Command, control, communications and intelligence 81

The German command system • The use of Kampfgruppen and the command system

Unit status 85

Lessons learned 91

Abbreviations and glossary 92

Bibliography 93

Key to vehicle silhouette identification 94

Index 95

© Osprey Publishing • www.ospreypublishing.com

 Introduction

Following the failure of the Kursk offensive, the German Army fought on the
defensive, first on the Eastern Front, then in Italy and on the Western Front. The
nature of warfare had changed again after the victorious years, and the German
Army adapted to meet the new requirements. Even the Panzerwaffe, the German
armoured force that once spearheaded the blitzkrieg campaigns, changed its
methods of operation. In turn, the role of the Panzer Divisions changed, as they
found themselves called upon to fight defensive battles. No longer were there
armoured thrusts into enemy territory; those that did happen served only to
relieve encircled units and allow them to escape. Tank battles were now mainly
fought to prevent the enemy from breaking through the German defence lines.
Furthermore, since late 1943 the Panzer Divisions of the Heer (Army) no longer
represented the sole component of the Panzerwaffe; new Waffen-SS Panzer
Divisions were upgraded or raised, while small armoured units, like the heavy
Tiger tank battalions, acquired greater importance.
Yet the Panzer Divisions still made up the bulk of the Panzerwaffe;
understrength, outnumbered and often fighting alone for months on end, they
faced the Soviet offensives on the Eastern Front and later spearheaded the
German forces both in Italy and in Normandy, eventually matching the role of
their Waffen-SS counterparts. Both served as ‘firefighters’ on the most
threatened areas of the front, as did other Panzer units such as the Tiger tank
battalions. Somehow, the role played by Army Panzer Divisions was less
glamorous and often comprised involvement in unknown battles fought in
unknown places. However, while the Waffen-SS could be called in at given
places and times, the Army Panzer Divisions still formed the backbone of the
German armed forces, despite their progressive weakening. In fact the overall
reorganization of the Panzerwaffe, started before Kursk, failed to restore their
overall potential and instead merely produced some high-quality divisions.
Although a lack of balance was not new in the Panzer Divisions, a new factor
further weakened their combat effectiveness: the dispersion of forces.
A PzKpfw V Panther Ausf D seeks On paper, the 1943 and 1944 Panzer Divisions were stronger than ever; new
cover somewhere in the battlefields tanks, a larger allotment of armoured vehicles for infantry and other units, self-
of Italy; once positioned, it will start propelled artillery and a new generation of tank hunters provided them with a
firing against enemy targets. The firepower no one would have dreamt of just a few years before. However, only
only Panther-equipped army unit
a few divisions ever came to be fully reorganized according to the intended
to see action in this theatre of
war was the I./Panzer Regiment 4, establishments, and many others fought relentlessly still equipped with the same
detached from the 13. to the 26. tanks, vehicles and weapons they had been using in early 1943. Lack of supplies,
Panzer Division. in particular fuel (which greatly hampered their combat capabilities), all
too often reduced them to the status of a small
Kampfgruppe, a battle group formed around the few
armoured and motorized units. Nonetheless, these
divisions were irreplaceable in their defensive roles and,
as a result, reorganization only affected the newly raised
ones and (to a certain extent) those that had been badly
mauled and refitted. Yet, in spite of their overall poor
shape, these divisions fought on all the fronts and still
proved capable of hitting back and spreading havoc
amongst their enemies. As the Ardennes offensive of
December 1944 proved, the Panzerwaffe was still able
to drive deep into enemy-held territory. However, this
4 time the enemy had changed.

© Osprey Publishing • www.ospreypublishing.com

Combat mission

Although the German Army was forced onto the defensive in the summer of
1943, its leading principles of warfare remained unchanged; attack, or
counterattack, was still the best way of achieving results against enemy forces.
Facing more enemies and fighting on several fronts simultaneously, the
German Army found in the Panzer Divisions a suitable instrument for a
makeshift solution: counterattacking enemy strikes when and where these
took place. Until 1944 German doctrine envisaged a linear defence based on a
main defence line, to be held at all costs regardless of enemy breakthroughs.
Mobile forces, above all the Panzer Divisions, were then ordered to
counterattack, surround and eventually destroy the enemy forces and restore
the situation. Such a system, which worked well on the Eastern Front until
mid-1944, had several shortcomings, however; in order to face enemy attacks
which threatened if not all at least good portions of the front, the Panzer
Divisions had to be scattered all along it because the lack of reserves and
supplies did not permit the switching of units fast enough to face the
impending crisis. This led to the exact opposite of one of the leading
principles of German warfare: the concentration of forces. As a result, the
Germans could still deal locally with enemy breakthroughs and restore the
situation (though often with severe losses), but they were no longer able to The crew of a PzKpfw IV Ausf H
regain the initiative and launch any major offensives. looking out for the approaching
Eventually, a lack of strength and resources led to the defence in depth enemy; the tank features all the late-
war improvements, which include
concept. Since the enemy relied heavily on firepower, the forward defensive
hull and turret skirts (Schürzen)
line was thinned down and comprised only a series of outposts intended to and the anti-magnetic mine paste
slow down the enemy attacking forces, while a main line of resistance was on the hull, known as ‘Zimmerit’.
built in the rear. When the enemy attack came, the Panzer Divisions had to The low-visibility numbers are
contain their drives and eventually counterattack following the usual also worth noting.
guidelines. Once again that led to a dispersion of forces and, as a result, no
major offensive was possible – as witnessed in Normandy. The debate
between Rommel and Von Rundstedt about the deployment of the Panzer
Divisions clearly reflects the situation; Rommel fully acknowledged Allied air
superiority and its impact on the Panzer Divisions’ operational capabilities,
while Von Rundstedt was seeking to regain the initiative through the
concentration of forces and a major offensive. Worth noting, both men were
right in their points of view; as the Ardennes offensive of December 1944
was to show, only a concentration of forces that managed to seek out a
Schwerpunkt – a ‘central gravity point’ – against the weaker enemy positions
could generate a breakthrough that might eventually lead to the surrounding
and destruction of enemy forces and the regaining of the initiative.
The fact that the Ardennes offensive took advantage of bad weather to deny
enemy air superiority does, on the other hand, show that Rommel was also
right. In fact, during the closing years of the war the Panzerwaffe was compelled
to wage war in a way for which it was not suited – i.e. a defensive war which
mainly sought to gain time or to hold ground rather than to seek out a decisive
victory on the field through manoeuvre. The Ardennes offensive was a doomed
one, but it was the last attempt made to try to put into practice the lessons of
German armoured warfare doctrine. When this was done, the odds were so
unfavourable that success never came within reach of the Panzer Divisions, even
though they proved again that they were still capable of demonstrating mastery
of the battlefield.
5

© Osprey Publishing • www.ospreypublishing.com

 Doctrine and training

There was no major change in German armour doctrine

in the closing years of the war, only adaptations to the
terrain and to new weapons and equipment. In 1943
the Panzer Regiment, or sometimes the Panzer
Abteilung, was the leading unit around which the main
Kampfgruppe (combat group) was built. In the closing
stages of the war many divisions were able to create
fully armoured Kampfgruppen that included elements
of Panzer, Panzergrenadier, Pionier, Panzerjäger,
artillery and other support units. It was employed like a
miniature Panzer Division, following the same rules of
armoured doctrine; Panzer units led the way, breaking
through the enemy defences and were closely
supported by self-propelled Artillerie, while follow-up
Close cooperation between the infantry, supported by Panzerjäger and Pionier units, secured the area and
Panzers and the Panzergrenadiers defended the flanks of the armoured drive. Although it remained in use until
increased towards the end of the 1945, this doctrine underwent several changes caused by new factors
war. The armament of this group
encountered on the battle front.
of Panzergrenadiers includes the
new assault rifle Sturmgewehr 44, First there were the new tactics of the Red Army, based on the creation of an
which indicates that they are part anti-tank defence screen to meet German armoured counterattacks; since
of a company’s Sturm Zug in action Panzer units could no longer afford huge losses, the infantry was now required
during the closing months of war. to deal directly with the anti-tank screen while the Panzers manoeuvred against
the enemy spearheads, often with the support of the Panzer Aufklärungs
Abteilung. This was now rarely used in its traditional reconnaissance role and,
because of its organization, was instead used as a regular combat unit. Another
decisive factor was Allied air superiority and firepower, which had a
considerable impact on German armoured doctrine. While on the Eastern
Front it was still possible to group large numbers of forces, in Italy (where Allied
naval support was so deadly) and in North-West Europe this was no longer
possible, since these large groups would have fallen prey to enemy air forces
and artillery. Therefore, the Panzer units no longer spearheaded the leading
Kampfgruppe but were rather split into smaller, company-sized units and used
to support the Panzergrenadier and the Panzer Aufklärungs units. The
Kampfgruppe became smaller, and doctrine returned to infiltration tactics,
aimed at achieving modest breakthroughs followed by swift advances. The
most logical consequence of this change was smaller, no longer decisive,
breakthroughs which had little impact.

Counterattacking enemy
breakthroughs, which included
Allied beachheads both in Italy
and in Normandy, was one of the
main tasks of the Panzer Divisions.
Here a Sturmgeschütz III, which
could equip both the Panzer and
the Panzerjäger Abteilung, is moving
6 past a destroyed Sherman tank.

© Osprey Publishing • www.ospreypublishing.com

 From 1943 onwards, both the Panzerkorps
(armoured corps) and the Panzerarmee (armoured
army) were armoured only in name. They turned into
a mixture of armoured and non-armoured units
which, though well-suited to defence, lacked the
necessary make-up for major offensive actions. The
need for closer cooperation between the Panzer
Divisions and the others, imposed by defensive needs,
added further limitations to the former. The battle for
Normandy offers a perfect example; in spite of the
concentration of armoured forces achieved in the area,
hardly any Panzer Divisions were grouped together to
launch a major counterattack; indeed, since the lack of
forces led to their deployment on the front line, they
were denied the advantages of mobility and speed.
Though still quite effective in defence, as the battles
on the German border showed, the Panzer Divisions
had clearly lost the edge.
This was despite their increased firepower, which was
greater than ever. The new Panther tank, though slow to
be delivered, provided Panzer units for the first time
with a tank that greatly outmatched its Allied
counterparts and dealt with the Soviet equivalents on a
even footing. Even anti-tank defences were improved
thanks to new anti-tank weapons like the Panzerschreck
and the Panzerfaust and to the new Jagdpanzer, which replaced the old Although the last models were
Panzerjäger with an all-round, heavily armoured gun carrier. The latter produced in August 1943, the
eventually brought about changes in German doctrine; turretless, self-propelled PzKpfw III remained in use (though
not with frontline units) until the end
gun carriers like the Sturmgeschütz and the new Jagdpanzer became offensive
of the war. Here we see a trainload
weapons, though they now required closer cooperation with the infantry. of PzKpfw IV Ausf G tanks, including
However, due to enemy air superiority movement was now mainly confined to some PzKpfw III Ausf M; with the
night-time or favourable weather, and anti-aircraft artillery (often self-propelled) November 1943 establishment,
was also required at every level – the presence of which further limited the these were still part of the
divisions’ speed and mobility. Panzer Abteilung’s Stabskompanie.
Last but not least, during the last two years of war the Panzer Divisions also
faced a major training crisis. The heavy personnel losses of the previous years, A Panzergrenadier veteran
plus those suffered during the summer of 1944, could only partly be made good Oberfeldwebel (technical sergeant)
using newly recruited personnel, and when this did happen divisional and unit in a training camp in Germany, in
commanders discovered that replacements needed a lot of additional training a staged propaganda photo. Other
before joining their experienced, battle-hardened comrades in arms. The Feld than a ‘wound’ badge, he displays
the iron crosses first and second
Ersatz Bataillon (the divisional field replacement battalion) allowed replacements
class, the infantry assault badge
to complete their training along with personnel from the division; this not only and the close combat badge. Such
refined their skills (for example in the use of heavy weapons and of artillery), but experienced men were essential for
also helped the recruits to acquaint themselves with battlefront conditions. training and leading the new recruits.
Technical training required by Panzer, Aufklärungs and Pionier units, on the
other hand, meant that these units were sent back to Germany to refit and
reorganize. Moreover, in 1944–45 the Germans faced serious problems due to a
shortage of trained officers and non-commissioned officers, which could only be
made good by promoting from the ranks. The situation became particularly
troublesome during the last, major reorganization in autumn 1944, in spite
of the overall, dramatic, reduction in strength imposed by the new tables
of organization.
All these problems notwithstanding, it is worth noting that even during the
difficult last year of war the Panzer Divisions performed well, often better than
their Waffen-SS counterparts, and – as the Ardennes offensive was eventually
to show – they still could achieve some decisive results.
7

© Osprey Publishing • www.ospreypublishing.com

 Unit organization

From 1943 to 1944

An order of the German Army High Command of 24 September 1943
introduced a common organization structure for all the Panzer Divisions (with
the exception of 21. Panzer and Panzer Division Norwegen) which, minor
variations apart, matched that already introduced in June for the Panzer
Divisions taking part in Operation Zitadelle. It was in practice a mere
acknowledgement of the status quo, since no overall reorganization was
possible due to the lack of men and equipment; nonetheless, on 1 November
1943 a new series of war establishment charts (KStN) were issued for the ‘1943
Panzer Division’ organization. This was to have a total strength of 16,385 all
ranks (439 officers, 89 officials, 3,781 NCOs, 12,076 ORs – 961 of which could
be replaced by foreign auxiliaries, the Hiwi – plus 800 replacements in the
Feldersatz Bataillon) armed with 10,808 rifles, 1,818 MPs, 695 light and 103
heavy MGs, 36 medium and 14 heavy mortars, 25 anti-tank guns plus 174
Panzerschreck, 25 field and four infantry guns and howitzers, eight 88mm and
38 20mm anti-aircraft guns (self-propelled ones not included), 771 cars, 1,946
lorries, 226 non-armoured half-tracks, 662 motorcycles, 215 tanks (of which 99

Basic organization of the Panzer Division type 1943 and 1944

Panzer Division

Begleit Komp

Panzer
Regiment Pz Gren Pz Gren
Regiment Regiment

I. Pz II. Pz
Abt Abt I. Btl II. Btl I. Btl II. Btl

Pz Aufkl Pz Jäg Pz Artillerie

Abt Abt Regiment HFA Abt

Pz Pionier PzDiv I. Abt II. Abt III. Abt

Bataillon Nachr Abt

Vers Tr FE Btl

© Osprey Publishing • www.ospreypublishing.com

were Panther tanks and 98 PzKpfw IV), 45 Panzerjäger and 405 AFVs, including
37 self-propelled guns of all types, 39 half-track mounted Flak guns, almost 300
armoured half-tracks (233 SdKfz 251 and 64 SdKfz 250) and 32 armoured cars.
In practice not a single division in the field matched its paper establishment,
chiefly because most of the PzKpfw V Panther-equipped Panzer Abteilungen
were still forming.
Also foreseeing the availability of new weapons, on 1 April 1944 the new
‘1944 Panzer Division’ organization was thus introduced, though the KStN
were still being issued in July. Actually, between April and May 1944 several
Panzer Divisions were meant to be reorganized, but only a few of them actually
completed the process before being committed to the front. On 3 August 1944
all divisions (except 21. Panzer and Panzer Lehr Division) were ordered to begin
reorganizing using available equipment and personnel, while reporting on
their status to allow for the issue of needed weapons and replacements. Overall
organization of the ‘1944 Panzer Division’ still matched that of the 1943
version, and changes mainly took place at regimental and battalion level. First
and foremost the new ‘freie Gliederung’ (free organization) was introduced,
which saw the creation for every battalion of a supply company (Versorgungs
Kompanie) from the supply units and trains of every company. Since the
commander of the Versorgungs Kompanie was also deputy battalion
commander, the new organization enabled the latter to focus exclusively on
combat duties, while the former took exclusive care of any logistical and
administrative needs.
Other changes affected overall strength and equipment; most notably,
infantry units lost their many anti-tank platoons, while heavy mortar and
flamethrower platoons were introduced. The overall strength of the 1944 Panzer
Division was 14,053 (411 officers, 85 Beamten, 3,219 NCOs, 10,338 ORs with 686
Hiwi plus 800 replacements) with 9,103 rifles, 1,637 MPs, 768 light and 98 heavy
MGs, 16 medium and 18 heavy mortars, 13 anti-tank guns and 9 Panzerschreck,
25 field guns and howitzers, eight 88mm, 36 20mm and six 37mm Flak guns, 303
cars, 852 lorries, 95 half-tracks and 171 motorcycles plus 179 tanks (79 Panther,
81 PzKpfw IV, eight Flakpanzer and 11 command and recovery tanks), 31
Panzerjäger and 372 AFVs, including 16 half-track mounted Flak guns, 36 gun
carriers, about 300 armoured half-tracks (245 SdKfz 251 and 59 SdKfz 250) and
16 armoured cars, all heavy. Even such a noticeable reduction failed to close the
growing gap between availability and need, further exacerbated by the heavy
losses suffered during the summer, and the overall lack of manpower and
equipment. Therefore, on 1 November 1944 new KStN were issued leading to the
late 1944 Panzer Division organization, which only affected Panzer and
Panzergrenadier regiments plus the Panzer Aufklärungs and Panzerjäger
Abteilung. The overall strength now dropped to 13,067 (416 officers, 77
Beamten, 2,964 NCOs, 9,610 ORs, 629 Hiwi, 800 replacements) with 8,719 rifles
and 270 Sturmgewehr 44, 1,527 MPs, 574 light and 32 heavy MGs, 12 medium

A PzKpfw V Panther Ausf D on

the battlefield. Many considered
it to be not only Germany’s best
tank, but also one of the best in
the world. This version weighed
43 tons, had a maximum speed of
45 kilometres per hour (25/30 off-
road) and a range of 200 kilometres
on road (100 off-road); its armour
was on average 80mm thick. 9

© Osprey Publishing • www.ospreypublishing.com

 Table 1: 1943 Panzer Division
Personnel Weapons
LMG Mortars
Offs Beamten NCOs ORs (Hiwi) Rifles MPs (HMG) (heavy)
Divisions Kommando (with Divisions Kartenstelle and Begleit Kompanie)
25 10 68 257 (17) 228 22 15 (2) 2
(+)4 x SdKfz 10/4
Panzer Regiment
63 9 880 1,466 (170) 1,004 296 46
(+) 6 x SdKfz 7/1 / 6 x SdKfz 251/7 / 4 x SdKfz 251/8
(1 x PzKpfw III / 2 x PzKpfw III PzBefh / 7 x SdKfz 141 (Pz III Flamm) / 99 x Panther / 98 x PzKpfw IV)
Panzergrenadier Regiment (gepanzert, with 9–11. Kompanie)
61 7 556 1,997 (84) 1,599 438 149 (35) 8 (4)
(+) 6 x SdKfz 138/1 / 37 x SdKfz 251/1 / 6 x SdKfz 251/2 / 20 x SdKfz 251/3 / 5 x SdKfz 251/4 / 6 x SdKfz 251/7 /
1 x SdKfz 251/8 / 12 x SdKfz 251/9 / 4 x SdKfz 251/10 / 7 x SdKfz 251/11 / 6 x SdKfz 251/16 / 22 x SdKfz 251/17 / 12 x SdKfz 10/4
Panzergrenadier Regiment (motorisiert, with 9–11. Kompanie)
57 6 503 1,947 (87) 1,585 281 149 (44) 14 (8)
(+) 6 x SdKfz 138/1 / 2 x ACs SdKfz 261 / 3 x SdKfz 251/1 / 1 x SdKfz 251/3 / 1 x SdKfz 251/11 / 12 x SdKfz 10/4
Panzer Aufklärungs Abteilung
26 3 279 765 (34) 622 189 49 (4) 2
(+) 12 x SdKfz 222 / 6 x SdKfz 223 / 3 x SdKfz 231 / 3 x SdKfz 232 / 6 x SdKfz 233 / 22 x SdKfz 250/1 /
2 x SdKfz 250/3 / 9 x SdKfz 250/5 / 4 x SdKfz 250/7 / 3 x SdKfz 250/8 / 16 x SdKfz 250/9 / 6 x SdKfz 251/1 /
11 x SdKfz 251/3 / 7 x SdKfz 251/7 / 1 x SdKfz 251/8 / 6 x SdKfz 251/9 / 3 x SdKfz 251/11 / 4 x SdKfz 251/17
Panzerjäger Abteilung
18 3 207 422 (43) 330 75 4
(+) 3 x SdKfz 7/1 / 1x SdKfz 251/8 45 x Panzerjäger (Marder II / III)
Panzer Artillerie Regiment
59 11 392 1,251 (77) 1,302 193 53

(*) 6 x PzBefh III (+) 3 x SdKfz 250/2 / 5 x SdKfz 250/5 / 12 (+ 4 ammunition carriers) x SdKfz 124 /
6 (+3 amm. Carriers) SdKfz 165 (*) 12 x le FH 105mm / 8 x schw FH 150mm / 4 x 10cm Kanone
Heeres Flak Artillerie Abteilung
21 3 174 679 (47) 767 58 22

(*) 4 x 20mm Flakvierling 38/1 SdKfz 7/1 (guns: 8 x 88mm Flak 36/37 / 24 x 20mm Flak 38 towed)
Panzer Pionier Bataillon
23 5 157 1,045 (74) 897 111 69 (6) 4
(+) 1 x SdKfz 251/1 / 2 x SdKfz 251/2 / 5 x SdKfz 251/3 / 23 x SdKfz 251/7 / 4 x SdKfz 251/10
Panzer Division Nachrichten Abteilung
13 3 103 396 (12) 444 51 13
(*) 2 x PzBefh III (+) 10 x SdKfz 251/3 / 6 x SdKfz 251/11 / 2 x SdKfz 251/19
Feldersatz Bataillon
17 1 91 64 (14) 73 86 50 (12) 6 (2)

2 x Flammenwerfer
Versorgungstruppen (with: Nachschub, Kraftfahr, Verwaltung, Sanitäts, Feldpost, Feldgendarmerie)
56 28 371 1,787 (302) 1,957 18 76
Note: the gun column only shows towed models, all others are included under AFVs.
10

© Osprey Publishing • www.ospreypublishing.com

 Another Random Document on
Scribd Without Any Related Topics
 NEGATIVE PHOTOTROPISM.

I shall next show the continuity of responsive phototropic effects, from the positive curvature to the
negative, through the intermediate phase of neutralisation. I have in the preceding paragraph described
an experiment where under a given intensity and duration of exposure the excitations of the proximal
and distal sides bring about neutralisation, the organ assuming a dia-phototropic position. If the
intensity or duration of the stimulating light be further increased, it is easy to see that while excitation
transmitted to the distal side is being increased, the excitatory contraction on the proximal side may, at
the same time, be decreased owing to fatigue brought on by over-stimulation.
In connection with this it should be borne in mind that the
pulvinus of Mimosa exhibits under continuous stimulation, a
fatigue relaxation instead of normal contraction. Similar effects
are known to take place in animal muscles. The effect of relatively
greater excitation will thus give rise to negative phototropic
curvature. The transverse conductivity of organs of diverse plants
will necessarily be different. The neutralisation and reversal into
negative will thus depend on three factors: the transverse
conductivity of the organ, the intensity, and duration of stimulus.
Neutralisation and reversal under increased intensity of light:
Experiment 127.—It is advisable to employ thin specimens (in
which the transverse distance is small) for the exhibition of
reversal effect. I took a hypocotyl of Sinapis nigra and subjected it
to unilateral action of light from a 16 candle-power incandescent
electric lamp placed at a distance of 10 cm. A maximum positive
curvature was induced in the course of 50 minutes. The intensity
of light was afterwards increased by bringing the lamp nearer to a
distance of 6 cm. This resulted in a process of neutralisation of
the preceding response; after an exposure of 70 minutes the
Fig. 121.—Positive and negative specimen assumed a dia-phototropic position in which it remained
phototropic responses of Oryza under in equilibrium. Sunlight was next applied, and in the further
continued unilateral stimulus of intense course of 30 minutes there was a pronounced reversal into
light from arc lamp. negative phototropic curvature.
Neutralisation and reversal under continuous stimulation:
Experiment 128.—In the last experiment the different changes in the response were brought about by
successive increase in the intensity of light. In the present experiment, very strong light was applied
from the beginning, and continuous record was taken of the change in the response. In order to reduce
the period of experiment I employed a mercury vapour lamp which emits the most effective violet and
ultra-violet rays. The specimen used was a seedling of the rice plant (Oryza sativa). The first effect of
light was a positive curvature which attained its maximum; after this there was a neutralisation in less
than six minutes after the application of light. The further continuation of light induced a pronounced
negative curvature (Fig. 121).
I shall in the next chapter give other instances which will show that all organs (pulvinated and
growing) possessed of power of transverse conduction, exhibit a transformation of response from
positive to negative under continued action of strong light.
Thus an identical organ, under different conditions of intensity and duration of stimulus, exhibits
positive phototropic, dia-phototropic, and negative phototropic curvatures, proving conclusively that the
three effects are not due to three distinct irritabilities. The responsive movements are, on the other
hand, traced to a fundamental excitatory reaction, remaining either localised or increasingly transmitted
to the distal side.
 NEGATIVE PHOTOTROPISM OF ROOTS.

From the analogy of opposite responses of shoot and root to stimulus of gravity, it was surmised
that the root would respond to light by a negative curvature. This was apparently confirmed by the
negative phototropic curvature of the root of Sinapis. The supposed analogy is however false; for while
the stimulus of gravity acts, in the case of root, only on a restricted area of the tip, the stimulus of light
is not necessarily restricted in the area of its action. That there is no true analogy between the action of
light and gravitation is seen from the fact that while gravitation induces in the root a movement
opposite to that in the stem, in the case of light, this is not always so; for though a few roots turn away
from light, others move towards the light.
As regards negative phototropic response of the root of Sinapis, it will be shown (p. 376) to be
brought about by algebraical summation of the effects of direct and indirect photic stimulus.

SUMMARY.

The normal positive phototropic curvature is modified by transverse conduction of true excitation to
the distal side of the organ.
The extent of neutralisation or reversal due to internal conduction of excitation from the proximal to
the distal side of the organ depends: (a) on the intensity of the incident stimulus, (b) on the
conductivity of the organ in a transverse direction, (c) on the thickness of the intervening tissue, and (d)
on the relative excitability of the distal as compared to the proximal side.
The dia-phototropic position is not one of indifference, but of balanced antagonistic reactions of two
opposite sides of the organ.
The supposition that direct sunlight is phototropically ineffective is unfounded. The response is fully
vigorous, but the first positive curvature may in certain cases be neutralised by the transmission of
excitation to the distal side.
Under light of strong intensity and long duration, the transmitted excitation to the distal side
neutralises, and finally reverses the positive into negative curvature.
The positive-phototropic, the dia-phototropic, and the negative phototropic curvatures are not due to
three distinct irritabilities but are brought about by a fundamental excitatory reaction remaining
localised or increasingly transmitted to the distal side.
[12] Jost—Ibid—p. 462.
[13] Jost—Ibid—p. 464.
 XXXI.—THE RELATION BETWEEN THE QUANTITY OF
LIGHT AND THE INDUCED PHOTOTROPIC CURVATURE
By

Sir J. C. Bose,

Assisted by

Surendra Chandra Das, M.A.

I shall in this chapter describe experiments in support of the important proposition that the intensity
of phototropic action is dependent on the quantity of incident light. The proportionality of the tropic
effect to the quantity of light will be found to hold good for the median range of stimulation; the
deviation from this proportionality at the two ends of the range of stimulation—the sub-minimal and
supramaximal—is, as we shall find, capable of explanation, and will be fully dealt with in the next
chapter.
The quantity of light incident on the responding organ depends: (1) on the intensity of light, (2) on
the angle of inclination or the directive angle,[14] and (3) on the duration of exposure. I shall give a
detailed account of the investigation relating to the individual effects of each of these factors on the
tropic reactions not merely in pulvinated but also in growing organs.

EFFECT OF INCREASING INTENSITY OF LIGHT ON TROPIC CURVATURE.

The intensity of light was increased in

successive experiments, in arithmetical
progression 1:2:3 by suitably diminishing the
distance between the plant and the source of
light, and the resulting tropic curvatures
recorded.
Effect of increasing intensity of light on the
pulvinus of Desmodium gyrans: Experiment
129.—The source of light was a 50 candle-
power incandescent lamp, and the duration of
exposure was 1 minute. The specimen
employed was a terminal leaflet of Desmodium
gyrans (Fig. 122) the pulvinus of which is very
sensitive to light. It is more convenient to
manipulate a cut specimen of the leaf, instead
of the whole plant. The petiole is placed in
Fig. 122.—Leaf of Desmodium gyrans, with the terminal water contained in a U-tube; the depressing
large, and two lateral small leaflets. These latter exhibit effect of wound passes off in the course of an
automatic pulsations. hour or so. Light of increasing intensity is
applied from above; this induces a contraction
of the upper half of the pulvinus, and the
resulting response is recorded by means of the Oscillating Recorder (Fig. 123).
 The first record was obtained under a given intensity, and the second, under an intensity twice as
great. The tropic effects are seen to increase with the intensity (Fig. 124). If the tropic curvature
increased proportionately to the intensity, the two responses should have been in the ratio of 1:2; the
actual ratio was however slightly greater, viz. 1:2·6. In this connection it will be shown in the next
chapter, that strict proportionality holds good only in the median range, and that the susceptibility for
excitation undergoes an increase at the beginning of the phototropic curve.

Fig. 123.—The Oscillating Recorder (From a photograph).

Fig. 124. Fig. 125.

 Fig. 124.—Tropic effect of
increasing intensity of light 1:2; on the
response of terminal leaflet of
Desmodium gyrans.
Fig. 125.—Tropic effect of
increasing intensity of light 1:2:3 on
growing organ (Crinum).

Effect of increasing intensity of light on the

tropic curvature of growing organs.—As the
tropic curvature is primarily due to the
retardation of growth induced by light at the
proximal side of the organ, it will be of interest
to recapitulate the results I obtained (p. 208) on
the effects of increasing intensity of light on
growth itself. The normal rate of growth of the
specimen in the dark was 0·47 µ per second;
this was reduced to 0·29 µ under an intensity of
one unit, to 0·17 µ under two, and to 0·10 µ under three units. Growth became arrested when the
intensity was raised to four units. Thus increasing intensity of light induces an increasing retardation of
growth at the proximal side of the organ. This aided by the effect of indirect stimulus at the distal side
brings about an increasing positive curvature.
Experiment 130.—The flower bud of Crinum was used for the experiment, the source of light being a
small arc lamp. The duration of exposure was one minute. Increasing intensity of light gave rise to
increasing positive curvatures (Fig. 125) in the ratio of 1:2·5:5 under increasing intensities which varied
as 1:2:3.

THE EFFECT OF INCREASING ANGLE.

The quantity of light which falls on an unit area of the responding organ varies as sin θ where θ is
the directive angle i.e. the angle made by the rays with the surface. Some allowance has to be made for
the amount of light reflected from the surface, this being greater at 45° than at 90°.

Fig. 126.—The Collimator.

Tropic response of pulvinus of Desmodium gyrans: Experiment 131.—For application of light at

various angles an incandescent electric lamp was mounted at one end of a brass tube, a collimating lens
being placed at the other (Fig. 126). The parallel beam of light from the collimator could be sent at
various angles by rotating the collimator tube round an axis at right angles to the tube. The specimen
employed was the terminal leaflet of Desmodium gyrans; light was applied for a minute in the two
successive experiments for the two angles of 45° and 90°. The record (Fig. 127) shows that the
phototropic effect increases with the directive angle. In the present case the ratio of the two effects is
1·6:1, which is not very different from the ratio sin 90°⁄sin 45° = 1·4.

Fig. 127. Fig. 128.

Fig. 127.—Effect of angle of inclination of light on the tropic curvature of pulvinus. The first response is to light at 45° and
to 90°. (Desmodium gyrans).
Fig. 128.—Series of tropic curvatures of growing bud of Crinum to alternate stimulation by light at 45° and 90°.

Tropic response of growing organs: Experiment 132.—Similar experiment was carried out with the
flower bud of Crinum, held vertical. Light was applied alternately at 45° and 90°, in two successive
series. The object of this was to make due allowance of possible variation of excitability of the organ
during the course of the experiment. I have explained (p. 147), how the excitability of a tissue in a
condition slightly below par, is increased by the action of previous stimulation. Series of responses
obtained under alternate stimulations at 45° and 90° enable us to find out, whether any variation of
excitability occurred during the course of the experiment and make allowance for it. The records show
that stimulation did enhance the excitability of the organ to a small extent. Thus the first stimulation at
45° induced an amplitude of response of 5 mm.; the second stimulation at 45° i.e. the third response of
the series, induced a slightly larger response 7 mm. in amplitude. Similarly the two responses at 90°
gave an amplitude of 9 mm. and 11 mm. respectively (Fig. 128). Taking the mean value of each pair,
the ratio of tropic effects for 90° and 45° is = 10⁄6 = 1·7 nearly.

EFFECT OF DURATION OF EXPOSURE.

Experiment 133.—The specimen employed for the experiment was a flower bud of Crinum in a
slightly sub-tonic condition. Successive responses exhibited on this account, a preliminary negative[15]
before the normal positive curvature. The successive durations of exposure were for 1, 2, and 3
minutes. The amplitudes of responses (Fig. 129) are in the ratio of 1:2·5:5.
 Fig. 129.—Effect of increasing duration of exposure 1:2:3 minutes, on phototropic
curvature. Note preliminary negative response. (Crinum).

We may now recapitulate the tropic effects of light of increasing intensity, directive angle, and
duration of exposure. It has been shown that the tropic effect is enhanced under increasing intensity of
light; it is also increased with the angle increasing from grazing to perpendicular incidence. And finally,
the tropic effect is enhanced with the duration of exposure. Taking into consideration the effects of
these different factors we arrive at the conclusion that phototropic effect increases with the quantity of
incident light. It will be shown in the next chapter that strict proportionality of cause and effect holds
good in the median range of stimulation, and the slight deviation from this, above and below the
median range, is due to the fact that susceptibility for excitation is low at these two regions.

SUMMARY.

Increasing intensity of light induces increasing tropic curvature.

Tropic curvature increases with the directive angle, the effect being approximately proportional to sin
θ, where θ is the angle made by the rays with the responding surface.
Tropic curvature also increases with the duration of exposure.
The intensity of induced tropic effect is determined by the quantity of incident light.
[14] The directive angle θ is the angle of inclination of the rays of light to the responding
surface. The angle θ is complementary to the angle of incidence i in optics. Sin θ = Cos i.
[15] An explanation of this preliminary effect will be found in the next chapter.
 XXXII.—THE PHOTOTROPIC CURVE AND ITS
CHARACTERISTICS
By

Sir J. C. Bose.
When a plant organ is subjected to the continued action of unilateral stimulus of light, it exhibits
increasing tropic curvature, which in certain cases reaches a limit; in other instances a reversal takes
place, seen in neutralisation, or in the conversion of the positive into negative curvature. I shall in this
chapter enter into a detailed study of the phototropic curve, and determine its characteristics.
As the vague terminology at present in use has been the source of much confusion, it is necessary
here to define clearly the various terms which will be employed in this investigation. It is first of all
necessary to distinguish between cause and effect, between external stimulus and the excitation
induced by it. As regards stimulus itself I have shown elsewhere[16] that its effective intensity becomes
summated by repetition. This was demonstrated by the two following typical experiments carried out
with the pulvinus of Mimosa.
(1) The intensity of a single electric shock of intensity of 0·5 unit was found to be ineffective in
inducing excitation; but it became effective on being repeated four times in rapid succession.
(2) The same specimen was next subjected to a feebler stimulus of intensity of 0·1 unit, and it
required a repetition of 20 times for the stimulus to become effective.
The total stimulus in the first case was 0·5 × 4 = 2, and this was found to be the same as 0·1 × 20
= 2 in the second case. Thus the intensity of stimulus is increased by repetition; in the limiting case
where the interval between successive stimulus is zero, the stimulus becomes continuous. Bearing in
mind the additive effects of stimulus we see that its effective intensity increases with the duration of
application. This important conclusion found independent support from the results of Experiment 133
given in the last chapter.
We shall now take up the general question of the characteristics of the phototropic curve, which
gives the relation between increasing stimulus and the resulting excitation. As regards stimulus we
found that its effectiveness increases with the duration of application. The induced excitation in growing
organs may be measured by concomitant retardation of growth caused by stimulus. In the excitation
curves which will be presently given, the abscissae represent increasing stimulus and ordinates the
resulting excitation. This excitation curve may be obtained by making the plant record on a moving
plate its retardation of growth by means of the High Magnification Crescograph. I reproduce below two
records of the effects of continuous photic and electric stimulation. The ordinate of the 'excitation curve'
(Fig. 130) exhibits increasing incipient contraction (retardation of growth) culminating in an arrest of
growth; the abscissa represents increasing stimulus consequent on increased duration of application.
The record shows that the incipient contraction is slight at the first stage; it increases rapidly in the
second stage; finally, it declines and reaches a limit. The excitatory reaction is thus not constant
throughout the entire curve of excitation, but undergoes very definite and characteristic changes. We
shall find similar characteristics in the phototropic curves under unilateral stimulus which will be given
presently. The explanation of the similarity is found in the fact that the tropic curvature is also due to
incipient contraction or retardation of the rate of growth, which remains confined to the directly
stimulated proximal side of the organ.
 Fig. 130.—Effects of continuous (a) electric, and (b) photic stimulation on rate of
growth. Abscissa represents duration of application of stimulus. Note induced
retardation, and arrest of growth.

For facility of explanation of what follows, I shall have to use a new and necessary term,
susceptibility, to indicate the relation of cause and effect, of stimulus and resulting excitation.
Susceptibility is thus = Excitation⁄Stimulus. Different organs of plants exhibit unequal susceptibilities; some
undergo excitation under feeble stimulus, while others require more intense stimulus to induce
excitation. But even in an identical organ the susceptibility undergoes, as we have seen, a characteristic
variation, being feeble at the beginning of the excitation curve, considerable in the middle, and
becoming feeble once more towards the end of the curve. The most difficult problem that faces us is an
explanation of this characteristic difference in different parts of the tropic curve.

GENERAL CONSIDERATIONS.

Before entering into the fuller consideration of the subject, it will be helpful to form some mental
picture of the phenomena of excitation, however inadequate it may be. The excitation is admitted to be
due to the molecular upset induced by the shock of stimulus[17]; the increased excitation results from
increasing molecular upset brought on by enhanced stimulus. The condition of molecular upset or
excitation may be detected from the record of any one of the several concomitant changes, such as the
change of form, (contraction or expansion) or change of electric condition (galvanometric negativity or
positivity). These means of investigation are not in principle different from those we employ in the
detection of molecular distortion in inorganic matter under increasing intensities of an external force.

THE CHARACTERISTIC CURVE.

Thus the molecular upset and rearrangement, in a magnetic substance under increasing magnetising
force are inferred from the curve obtained by means of appropriate magnetometric or galvanometric
methods. I reproduce the characteristic curve of iron (Fig. 131) in which the abscissa represents
increasing magnetising force, and the ordinate, the induced magnetisation. This characteristic curve,
giving the relation of cause and effect, will be found to be highly suggestive as regards the similar
characteristic curve which gives the relation between increasing stimulus and the resulting enhanced
tropic effect in vegetable tissues. The parallelism will be found to be very striking.
Inspection of figure 131 shows that, broadly speaking, the curve of magnetisation may be divided
into four parts. In the first part, under feeble magnetic force, the slope of the curve is very small; later,
in the second part, as the force increases, the curve becomes very steep; in the third part the slope of
the curve remains fairly constant; and finally in the fourth part, the curve rounds off and the rate of
ascent again becomes very small. The susceptibility for induced magnetisation is thus very feeble at the
beginning; under increasing force, in the second stage, the susceptibility becomes greatly enhanced; in
the third stage, the susceptibility remains approximately constant; and in the fourth stage it becomes
diminished. We shall presently find that the susceptibility for excitation also undergoes a similar
variation at the four different stages of stimulation.

CHARACTERISTICS OF SIMPLE PHOTOTROPIC CURVE.

I have shown (Fig. 130) the relation between the stimulus and the resulting excitation, the latter
being determined from the diminution of the rate of growth. Under unilateral action of light, the
excitatory contraction gives rise to tropic curvature. We may thus obtain the characteristic excitation
curve, by making the plant organ record its tropic movement under continuous action of light applied on
one side of the organ.

Fig. 131. Fig. 132.

Fig. 131.—Characteristic curve of iron under increasing magnetising force. (After Ewing).
Fig. 132.—Simple characteristic curve of phototropic reaction. Excitation increases slowly in the first part
and rapidly in the second; it is uniform in the third, and undergoes decline in the fourth part (Erythrina
indica).

Experiment 134.—I give below the characteristic curve of excitation (Fig. 132) of the pulvinus of
Erythrina indica, traced by the plant itself, and exactly reproduced by photomechanical process. A
parallel beam of light from a Nernst lamp was thrown on the upper leaf of the pulvinus, and the
increasing positive curvature was recorded on a smoked glass plate which was moved at an uniform
rate. The successive dots are at intervals of 20 seconds; the horizontal distances between successive
dots are equal, and represent equal increments of stimulus; the vertical distances between successive
dots represent the corresponding increments of excitation. The gradient at any point of the curve—
increment of excitation divided by increment of stimulus—gives the susceptibility for excitation at that
point. The following table will show how the susceptibility for excitation undergoes variation through the
entire range of stimulus. The average susceptibility for each point has been calculated from the data
furnished by the curve.
 TABLE XXX.—SHOWING THE VARIATION OF SUSCEPTIBILITY FOR EXCITATION AT
DIFFERENT POINTS OF THE TROPIC CURVE.

Successive points in the Susceptibility for Successive points in the Susceptibility for
curve. excitation. curve. excitation.
1 0 14 6·6
2 0·187 15 4·4
3 0·44 16 2·5
4 0·625 17 1·87
5 0·875 18 1·5
6 1·25 19 1·12
7 1·87 20 0·937
8 3·12 21 0·75
9 5·0 22 0·562
10 6·25 23 0·375
11 8·75 24 0·25
12 8·87 25 0·187
13 8·12 26 0·062

The induced excitation is seen to be increased very gradually from the zero point of susceptibility,
known as the latent period at which no excitation takes place. In the second part of the excitation
curve, the rate of increase is vary rapid; the maximum rate is nearly reached at point 11 of the curve
and remains fairly constant for a time. This is the median range where equal increment of stimulus
induces equal increment of excitation. The susceptibility for excitation then falls rapidly, and increase of
stimulus induces no further increase of tropic curvature. The maximum tropic curvature was, in the
present case, reached in the course of nine minutes. The attainment of this maximum depends on the
excitability of the tissue, and the intensity of incident stimulus. The characteristics that have been
described are not confined to the phototropic curve but exhibited by tropic curves in general. Similar
characteristics have been found in the curve for electric stimulus (Fig. 130a), and will also be met with
in the curve for geotropic stimulus (Fig. 161).
I may here refer incidentally to the three types of responses exhibited by an organ to successive
stimuli of uniform intensity; these appear to correspond to the three different regions of tropic curve; in
the first stage, the plant exhibits a tendency to exhibit a 'staircase' increase of response; in the
intermediate stage, the response is uniform; and in the last stage, the responses show a 'fatigue'
decline.
For purpose of simplicity, I first selected the simple type of phototropic curve, where the specimen
employed was in a favourable tonic condition, and the stimulus was, from the beginning, above the
minimal. Transverse conduction, which induces neutralisation or reversal into negative, was moreover
absent in the specimen. I shall now take up the more complex cases: (1) where the condition of the
specimen is slightly sub-tonic, (2) where the stimulus is gradually increased from the sub-minimal, and
(3) where the specimen possesses the power of transverse conduction.

EFFECT OF SUB-MINIMAL STIMULUS.

It is unfortunate that the terms in general use for description of effective stimulus should be so very
indefinite. A stimulus which is just sufficient to evoke excitatory contraction is termed the minimal,
stimulus below the threshold being tacitly regarded as ineffective. The employment of sensitive
recorders has, however, enabled me to discover the important fact that stimulus below the minimal,
though ineffective in inducing excitatory contraction, is not below the threshold of perception. The plant
not merely perceives such stimulus, but also responds to it in a definite way, by expansion instead of
contraction. I shall designate the stimulus below the minimal, as the sub-minimal. There is a critical
point, which demarcates the sub-minimal stimulus with its expansive reaction from the minimal with its
responsive contraction.
The critical stimulus varies in different species of plants. Thus the same intensity of light which
induces a retardation of growth in one species of plants will enhance the rate of growth in another.
Again, the critical point will vary with the tonic level of the same organ; in an optimum condition of the
tissue, a relatively feeble stimulus will be sufficient to evoke excitatory contraction; the critical point is
therefore low for tissues in tonic condition which may be described as above par. In a sub-tonic
condition, on the other hand, strong and long continued stimulation will be necessary to induce the
excitatory reaction. The critical point is therefore high, for tissues in a condition below par. Stimulus
below the critical point will here induce the opposite physiological reaction, i.e., expansion. The physico-
chemical reactions underlying these opposite physiological responses have, for convenience, been
distinguished as the "A" and "D" change (pp. 143, 223). The assimilatory 'building up', A change, is
associated with an increase of potential energy of the system; the dissimilatory 'break down', D change,
on the other hand, is attended by a run-down of energy.
Stimulus was shown (p. 225) to give rise to both these reactions, though the A effect is, generally
speaking, masked by the predominant D effect. The "A" change is quicker in initiation, while the "D"
effect developes later; again the "A" effect under moderate stimulation may persist longer. Thus owing
to the difference in their time-relations the A effect is capable of being unmasked at the onset of
stimulus or on its sudden cessation. For the detection of the relatively feeble expansive A effect, a
special recorder is required which combines lightness with high power of magnification. The earlier
expansive reaction and acceleration of rate of growth, followed by normal retardation, are often found
in the response of growing organs. The corresponding effect of unilateral stimulation, even when direct,
is a transient expansion at the proximal side, inducing a convexity of that side and movement away
from stimulus (negative curvature); this is followed by contraction and concavity with normal positive
curvature. The interval between the A and D effects is increased with increasing sub-tonicity of the
specimen. But it nearly vanishes when the excitability of the specimen is high, and the two opposite
reactions succeed each other too quickly for the preliminary A reaction to become evident. It is probable
that in such a case the conflict between the two opposite reactions prolongs the latent period. But in
other instances a preliminary expansive response is found to herald the more pronounced contractile
response. Example of this is seen in figure 129 given in page 344.
The A effect was detected in the records referred to above by its earlier appearance. Its longer
persistence, after moderate stimulation, is also to be found on the cessation of moderate stimulation.
This was seen in the acceleration of growth which was the after-effect of stimulation (Figs. 104, 115).
The presence of two conflicting physiological reactions is also made evident on sudden cessation of long
continued stimulation. This particular phenomenon of "overshooting" will be more fully dealt with in a
subsequent chapter.
Owing to the difference in the time relations of the two opposing activities, A and D, a phase
difference often arises in their respective maxima. It is probably on this account that rhythmic tissues
originally at standstill, exhibit under continued stimulation a periodic up and down-movement, which
persists even on the cessation of the stimulus. The persistence of after-oscillation depends, moreover,
on the intensity and duration of previous stimulation.[18]
The facts given above cannot be explained by the prevalent theory that stimulus acts merely as a
releasing agent, to set free energy which had been previously stored up by the organism, like the pull of
a trigger causing explosion of a charged cartridge. It is true that in a highly excitable tissue, the
external work performed and the run down of energy are disproportionately greater than the energy of
stimulus that induces it. But in a sub-tonic tissue, stimulus induces an effect which is precisely the
opposite; instead of a depletion, there is an enhancement of potential energy of the system. Thus the
responding leaf instead of undergoing a fall becomes erected; growing organs similarly exhibit a
'building up' and an acceleration of rate of growth, in contrast with the usual 'break down' and
depression of the rate. It is obvious that these new facts relating to the action of stimulus necessitate a
theory more comprehensive and satisfactory than the one which has been in vogue.

THE COMPLETE PHOTOTROPIC CURVE.

I have explained the characteristics of the simple phototropic curve in which the tropic curvature, on
account of the favourable tonic condition and strong intensity of incident light, was positive from the
beginning, and in which the curvature reached a maximum beyond which there was no subsequent
reversal. If the intensity of the stimulus be feeble or moderate, the quantity of light incident on the
responding organ at the beginning may fall below the critical value, and thus act as a sub-minimal
stimulus. This induces as we have seen (p. 344) a negative tropic curvature; continued action of
stimulus, however, converts the preliminary negative into the usual positive. The preliminary negative
curvature may be detected by the use of a moderately sensitive recorder with a magnification of about
30 times. It is comparatively easy to obtain the preliminary negative response in specimens which are in
a slightly sub-tonic condition.
Semi-conducting tissues exhibit under continued stimulation, a neutralisation and reversal into
negative (p. 331). Since this reversal into negative usually takes place under prolonged exposure to
exceedingly strong light, it is difficult to obtain in a single curve all the different phases of
transformation. I have, however, been fortunate in obtaining a complete phototropic curve which
exhibits in a single specimen all the characteristic changes from a preliminary negative to positive and
subsequent reversal to negative. I shall describe two such typical curves obtained with the terminal
leaflet of Desmodium gyrans and the growing seedling of Zea Mays.
Complete phototropic curve of a pulvinated organ: Experiment 135.—A continuous record was taken
of the action of light of a 50 c. p. incandescent lamp, applied on the upper half of the pulvinus of the
terminal leaflet of Desmodium gyrans. This gave rise: (1) to a negative curvature (due to sub-minimal
stimulus) which lasted for 3 minutes. The curve then proceeded upwards, at first slowly, then rapidly; it
then rounded off, and reached a maximum positive value in the course of 18 minutes; after this the
curve underwent a reversal, and complete neutralisation occurred after a further period of 24 minutes
(Fig. 133). Beyond this the induced curvature is negative.

Fig. 133.—Complete phototropic curve given by pulvinated Eq. organ. Positive

curvature above, and negative curvature below the horizontal zero line. Preliminary
negative phase of response due to sub-minimal stimulus. The positive increases,
attains a maximum, and undergoes a reversal. Successive dots at intervals of 30
seconds. Abscissa represents duration of exposure and quantity of incident light.
(Terminal leaflet of Desmodium gyrans.)

Complete phototropic curve of growing organs: Experiment 136.—I obtained very similar effects by
subjecting the seedling of Zea Mays to unilateral light from an arc lamp for two hours. The characteristic
of this curve is similar to that given by the terminal leaflet of Desmodium gyrans. At the first stage, the
sub-minimal stimulation is seen to induce a negative curvature, transformed into positive after an
interval of 10 minutes. The maximum positive curvature is reached after 50 minutes, and neutralisation
completed in a further period of 43 minutes (Fig. 134). After this the response became transformed into
negative.

Fig. 134.—Complete phototropic curve of a growing organ (Zea Mays).

In a complete phototropic curve we may thus distinguish 4 distinct stages:—

(1) The stage of sub-minimal stimulation.
(2) The stage of increasing positive curvature culminating in a maximum.
(3) The stage of neutralisation.
(4) The stage of complete reversal into negative.
The curve thus crosses the zero line of the abscissa twice; the first crossing takes places upwards at
the critical point of stimulation which demarcates the sub-minimal from the minimal. The second
crossing downwards occurs beyond the point of complete neutralisation.
In a tissue in which transverse conductivity is absent, and the stimulus applied from the beginning is
above the minimal, the simple tropic curve is confined to the second stage (see Fig. 132).

WEBER'S LAW.

If we neglect the preliminary negative portion under sub-minimal stimulus, the curve of excitation
under increasing photic stimulation obeys what is known as Weber's law. This is equally true of modes
of stimulation other than that of light as is seen in figure 130 of the contractile effect of continued
electric stimulus on growth; the excitatory effect is also seen to reach a limit.
Weber's law is applicable for a limited range of stimulation. For the quantitative relation fails in the
region of sub-minimal stimulus, where the physiological reaction is qualitatively different, namely
expansion instead of contraction. This holds good even in the case of animal tissues, for here also my
recent experiments show that two opposite reactions—expansion and contraction—take place under
stimulus, and that a very feeble stimulus tends to induce expansion instead of contraction. The
responsive reaction of a kitten under gentle caressing strokes must be qualitatively different from that
of a blow. The psychological effects under the two treatments evidently differ qualitatively rather than
quantitatively.[19]

SUMMARY.
 The excitation curve exhibits a slow ascent in the first part; in the second part the gradient is steep,
indicating rapid rise in excitation; in the third part it is uniform; and in the last part the curve rounds off
and the rate of ascent becomes very small.
The susceptibility for excitation is feeble at the beginning; it increases very rapidly with increasing
stimulus; finally it undergoes a fall, increase of stimulus inducing no further enhancement of excitation.
In a complete phototropic curve the first part is negative; this is due to the physiological expansion
induced by sub-minimal stimulus. The curve then crosses the abscissa upwards, and the positive
curvature reaches a maximum. This is followed by neutralisation and reversal into negative; the curve
crosses the zero line and proceeds in the negative direction.
Weber's law is not applicable for the entire range of stimulation. The quantitative relation fails in the
region of sub-minimal stimulus, where the physiological reaction is qualitatively different.
[16] "Irritability of Plants"—p. 54.
[17] I shall use the term stimulus in preference to stimulation, for the latter is often taken in the
sense of the resulting excitation.
[18] "Plant Response"—p. 293, etc.
[19] "It has been argued by James that the feeling does not cause, but is caused by the bodily
expression.... Münsterberg concludes that the feeling of agreeableness is the mental
accompaniment and outcome of reflexly produced movements of extension, and disagreeableness
of the movement of flexion." Schāfer—Text Book of Physiology, Vol. II, p. 975 (1900).
 XXXIII.—THE TRANSMITTED EFFECT OF PHOTIC
STIMULATION
By

Sir J. C. Bose,

Assisted by

Jyotiprakash Sircar, M.B.

Plant organs exhibit, as we have already seen, a heliotropic curvature under direct stimulation. Still
more interesting is the transmitted effect of light giving rise to a curvature. Thus if the tip of the
seedling of wheat be exposed to light, the excitation is transmitted lower down into the region which
acts as the responding organ. Growth is very active in this particular zone, and the change of growth,
induced by the transmitted effect of stimulus, brings about a curvature by which the tip of the seedling
bends towards light. The seedling thus appears to be differentiated into three physiological zones
subserving three different functions. The tip is the perceptive zone, the intervening distance between
the tip and the growing region is the zone of conduction, and the growing region is the responsive zone.
These differentiations are shown in a striking manner by certain Paniceae, Setaria for example. In this
seedling the tapering sheathing leaf or cotyledon is about 5 mm. in length, and it is the upper part of
the cotyledon that is most sensitive to light. Below the sheathing leaf is a narrow length which will be
distinguished as the hypocotyl, and where growth is very active. The apex of the leaf perceives the
stimulus, and the effect is transmitted to the hypocotyl, which responds by becoming curved so that the
seedling bends towards light.
It is necessary here to make special reference to the confusion that arises from want of precision in
the use of the term stimulus, used indifferently to denote both the cause and the resulting effect. An
external agent, say light, causes certain excitatory change in the tissue, and we refer to the agent
which induces it, as the stimulus. Thus in the instance cited above, light is the stimulus, and it is the
stimulus-effect that is transmitted to a distance. But in physiological literature no distinction is made
between the stimulus and its effect, hence arises frequent use of the phrase 'transmission of stimulus'.
It is obvious that it is not light but its effect that is transmitted.
Such want of precision in the use of the term stimulus would not have seriously affected the truth
about the description of facts, had the transmitted effect been only of one kind. In a nerve-and-muscle
preparation, the velocity of transmission of excitation is so great, that it completely masks the positive
impulse (assuming the existence of such an impulse). The effect of indirect stimulation is, therefore, the
same as that of direct stimulation. Any indefiniteness in the use of the term stimulus for its transmitted
effect does not, in animal physiology, seriously militate against the observed facts. But lack of precision
in the employment of the term in plant physiology leads to hopeless confusion. For owing to the semi-
conducting nature of vegetable tissue, the transmitted effect is not of a definite sign, but may be
positive or negative; in the first case, the response is by expansion, in the latter, by contraction. Thus
the transmitted effect will be very different in the two cases, according as the intervening tissue is a
good or a bad conductor. These facts accentuate the urgent necessity of revision of our existing
terminology.
 I have shown that the effects of other forms of stimuli are also transmitted from the perceptive to
the responding region along the intervening path of conduction. Thus the petiole of Mimosa perceive
any form of stimulus applied to it, and the induced excitation is conducted to the distant pulvinus to
evoke the familiar responsive fall of the leaf. The pulvinus, moreover, perceives and responds to direct
stimulation. In a nerve-and-muscle preparation the responding muscle is alike perceptive and
responsive.
But in Setaria we meet with certain characteristics of reaction which are quite inexplicable. Thus if
"the seedling be illuminated on one side, a sharp heliotropic curving takes place at the apex of
hypocotyl. The curvature makes itself apparent only if the cotyledon be illuminated from one side
whether the hypocotyl be exposed to light or not. If the cotyledon be shaded and the light be permitted
to fall on one side of the hypocotyl, no heliotropic curving takes place. Hence we may conclude that it is
only the cotyledon that is sensitive to the light stimulus, and it is only the hypocotyl which can carry out
the movement. The excitation which the light effects in the cotyledon must be transmitted to the
hypocotyl and curvature takes place only from such a transmitted excitation. We have thus in this case a
definite organ for the perception of the stimulus of light, viz., the cotyledon, and as Rothert has shown,
it is more specially the apex of that organ that is the sensitive part: on the other hand, the motile
organ, the hypocotyl, is some distance away from the sensitive organ, and in it the power of perception
is entirely absent. From the behaviour of these organs we may draw the further conclusion that
perception and heliotropic excitation are two distinct phenomena, which depend on different properties
of the protoplasm and which are independent of each other.... We may, therefore, conclude from this
experiment that these two types of excitation are fundamentally distinct processes, for it is only after
indirect or transmitted and not after direct excitation that a reaction occurs in the case of the seedlings
of the Paniceae".[20]
The noteworthy deductions on the above facts are:—
(1) That the motile organ in Setaria is totally devoid of perception, since direct action of
light induces no effect.
(2) That perception and heliotropic excitation are two distinct phenomena, which depend
on different properties of the protoplasm, and which are independent of each other.
Though the conclusions thus arrived at appear to follow from the facts that have been observed, yet
it is difficult to accept the inference, that a responding organ should be totally devoid of the power of
perception, and that excitation and perception are to be regarded as dependent on different properties
of protoplasm. It therefore appeared necessary to re-investigate the subject of the perceptive power of
the cotyledon, and the responding characteristics of the hypocotyl.
The criterion employed for test of perception is the movement induced in response to stimulus. The
responsive mechanical movement is rendered possible only by the contractility of the organ, and
mechanical and anatomical facilities offered by it for unhampered movement. The petiole of Mimosa
when locally stimulated does not itself exhibit any movement. The fortunate circumstance of the
presence of a motile pulvinus in the neighbourhood enables us to recognise the perceptive power of the
petiole, since it transmits an impulse which causes the fall of the leaf. There is no motile pulvinus in
ordinary leaves, and stimulation of the petiole gives rise to no direct or transmitted motile reaction;
from this we are apt to draw the inference that the petiole of ordinary leaves are devoid of perception.
This conclusion is, however, erroneous, since under stimulus the petiole exhibits the electric response
characteristic of excitation. Moreover my electric investigations have shown that every living tissue not
only perceives but also responds to stimulation.[21] Hence considerable doubt may be entertained as
regards the supposed absence of perception in the hypocotyl of Setaria.
I shall in the present paper describe my investigations on the mechanical response of Setaria under
direct and indirect stimulation which will be given in the following order:—
 (1) The response to unilateral stimulation of the tip of the seedling.
(2) The response of growing hypocotyl to direct stimulation.
(3) Summated effects of direct and indirect stimulation.

EXPERIMENTAL ARRANGEMENTS.

The Recorder.—The pull exerted by the tropic curvature of the seedling is very feeble; it was
therefore necessary to construct a very light and nearly balanced recording lever. A long glass fibre is
supported by lateral pivots on jewel bearings. The seedling is attached to the short arm of the lever by
means of a cocoon thread. The recording plate oscillates to and fro once in a minute; the successive
dots give therefore the time relations of the responsive movement. The positive curvature towards light
is recorded as an up-curve, the negative curvature being represented by a down-curve.

Fig. 135.—Arrangement for local application of light to the tip and the growing
region. O, O', apertures on a metallic screen. Light is focussed by a lens on the tip,
and on the growing region at o, o'. Figure to the right shows front view of the shutter
resting on a pivot and worked by string, T.

Arrangement for local stimulation by light.—The device of placing tin foil caps on the tip employed by
some observers labours under the disadvantage, that it causes mechanical irritation of the sensitive tip.
The appliance seen in figure 135 is free from this objection and offers many advantages. A metallic
screen has two holes O and O'; these apertures are illuminated by a parallel beam of light from an arc
lamp. A lens focusses the light from O, on the hypocotyl, and that from O', on the tip of the cotyledon.
A rectangular pivoted shutter S, lies between the apertures O and O'. In the intermediate position of the
shutter, light acts on both the tip and the growing region. The shutter is tilted up by a pull on the thread
T, thus cutting off light from the growing region; release of the thread cuts off light from the tip. Thus
by proper manipulation of the shutter, the tip or the growing hypocotyl, or both of them, may be
subjected to the stimulus of light. The experiment was carried out in a dark room, special precaution
being taken that light was screened off from the plant except at points of localised stimulation.
 Fig. 136.—Response of seedling of Setaria to unilateral stimulation of the tip applied at
dotted arrow.
Note preliminary negative curvature reversed later into positive.

EFFECT OF LIGHT AT THE TIP OF THE ORGAN.

Experiment 137.—If the tip of the seedling of Setaria be illuminated on one side, it is found that a
positive curvature (i.e., towards light) is induced in the course of an hour or more. But in obtaining
record of the seedling by unilateral stimulation of the tip, I found that the immediate response was not
towards, but away from light (negative curvature). The latent period was about 30 seconds and the
negative movement continued to increase for 25 minutes (Fig. 136). This result, hitherto unsuspected,
is not so anomalous as would appear at first sight. Indirect stimulus, unilaterally applied, has been
shown to give rise to two impulses: a quicker positive and a slower excitatory negative. The former
induces a convexity on the same side, and a movement away from stimulus (negative curvature); the
excitatory negative, on the other hand, is conducted slowly and induces contraction and concavity, and
a movement towards the stimulus (positive curvature). In semi-conducting or non-conducting tissues,
the excitatory negative is weakened to extinction during transit, and the positive reaction with negative
curvature persists as the initial and final effect.
But in Setaria the excitatory negative impulse is transmitted along the parenchyma which is
moderately conducting; the speed of transmission of heliotropic excitation is, according to Pfeffer, one
or two mm. in five minutes or about 0·4 mm. per minute. Thus under the continued action of light, the
excitatory impulse will reach the growing region, and by its predominant reaction neutralise and reverse
the previous negative curvature.
Inspection of figure 136 shows that this is what actually took place; the intervening distance
between the tip of the cotyledon and the growing region in hypocotyl was about 20 mm., and the
beginning of reversal from negative to positive curvature occurred 29 minutes after application of light.
The velocity of transmission of excitatory impulse under strong light is thus 0·7 mm. per minute. The
positive curvature continued to increase for a very long time and became comparatively large. This is for
two reasons: (1) because the sensibility of the tip of the cotyledon is very great, and (2) because the
positive curvature induced by longitudinally transmitted excitation is not neutralised by transverse
conduction (see below).
 Fig. 137.—Effect of application of light to the growing hypocotyl at arrow induced
positive phototropic curvature followed by neutralisation. Application of indirect
stimulus at dotted arrow on the tip gave rise at first to negative, subsequently to
positive curvature. (Seedling of Setaria).

RESPONSE TO UNILATERAL STIMULUS IN THE GROWING REGION.

Experiment 138.—The growing region of the hypocotyl of Setaria is supposed to be totally devoid of
the power of perception. In order to subject the question to experimental test, I applied unilateral light
on the growing region of the same specimen, after it had recovered from the effect of previous
stimulation. The response now obtained was vigorous and was ab-initio positive. Direct stimulus has
thus induced the normal effect of contraction and concavity of the excited side. The belief that the
hypocotyl of Setaria is incapable of perceiving stimulus is thus without any foundation. The further
experiment which I shall presently describe will, however, offer an explanation of the prevailing error.
On continuing the action of unilateral light, the positive curvature after attaining a maximum in the
course of 15 minutes, underwent a diminution and final neutralisation (Fig. 137). On account of this
neutralisation the seedling became erect after an exposure of 30 minutes; in contrast with this is the
increasing positive curvature under unilateral illumination of the tip (Fig. 136) which continues for
several hours. The explanation of this neutralisation under direct stimulation of the growing region is
found in the fact that transverse conduction of excitation induces contraction at the distal side of the
organ and thus nullifies the positive curvature. The seeming absence of tropic effect under direct
stimulation is thus not due to want of perception, but to balanced antagonistic reactions on opposite
sides of the organ.

EFFECT OF SIMULTANEOUS STIMULATION OF THE TIP AND THE HYPOCOTYL.

Though stimulation of the hypocotyl results in neutralisation, yet the illumination of one side of the
organ including the tip and hypocotyl is found to give rise to positive curvature. This will be understood
from the following experiment.
After the neutralisation in the last experiment light was also applied to the tip from the right side at
the dotted arrow (Fig. 137). The record shows that this gave rise at first to a negative curvature (away
from light); under the continued action of light, however, the negative was subsequently reversed to a
positive curvature, towards light. Inspection of the curve shows another interesting fact. The positive
curvature induced by direct stimulation is very much less than that brought out by indirect stimulation.
This is due to two reasons: (1) the sensitiveness of the tip of the organ is, as is well known, greater
than that of the hypocotyl, (2) the positive curvature under direct stimulation cannot proceed very far,
since it is neutralised by transverse conduction of excitation.
It will be seen from the above that the illumination of the tip practically inhibits the neutralisation
and thus restores the normal positive curvature. The question now arises as to how this particular
inhibition is brought about.
 ALGEBRAICAL SUMMATION OF THE EFFECTS OF DIRECT AND INDIRECT STIMULATIONS.

An instance of inhibition, though of a different kind, was noticed in the response of the tendril of
Passiflora (p. 296); the under side of the organ is highly sensitive, while the upper side is almost
insensitive. Stimulation of the under side of the tendril induces a marked curvature, but simultaneous
stimulation of the diametrically opposite side inhibits the response. This neutralisation could not be due
to the antagonistic contraction of the upper side since the irritability of that side is very slight. I have
shown that the inhibition results from the two antagonistic reactions, contraction at the proximal side
due to direct stimulation and expansion caused by the positive impulse from the indirectly stimulated
distal side.
We have in the above an algebraical summation of the effects of direct and indirect stimulations. The
longitudinally transmitted effect of indirect stimulus in Setaria may, likewise, be summated with the
effect of direct stimulus. The phenomenon of algebraical summation is demonstrated in a very striking
and convincing manner in the following experiment, which I have been successful in devising.
Experiment 139.—I have explained, (Expt. 126) that unilateral application of stimulus of light on the
upper half of the responding pulvinus of Mimosa induces an up or positive curvature, followed by a
neutralisation and even a reversal into negative, the last two effects being brought about by transverse
conduction of excitation to the distal side. When the incident light is of moderate intensity, the
transmitted excitation only suffices to induce neutralisation without further reversal into negative; while
in this state of balanced neutralisation let us apply indirect stimulus by throwing light on the stem at a
point directly opposite to the leaf (Fig. 138).

Fig. 138.—(a) Diagrammatic representation of direct application of light (↓) on the

pulvinus and the indirect application on the stem (→) (b) Record of effect of direct
stimulus, positive curvature followed by neutralisation. Superposition of the positive
reaction of indirect stimulus induces erectile up-response followed by down movement
due to transmitted excitatory impulse (Mimosa).

Two different impulses are thus initiated from the effect of indirect stimulus. In the present case the
positive reached the responding pulvinus after 30 seconds and induced an erectile movement of the
leaf; the excitatory negative impulse reached the organ 4 minutes later and caused a rapid fall of the
leaf. The record (Fig. 138) shows further that the previous action of direct stimulus which brought about
neutralisation, does not interfere with the effects of indirect stimulus. The individual effects of direct and
indirect stimulus are practically independent of each other; hence their joint effects exhibit algebraical
summation.
We are now in a position to have a complete understanding of the characteristic response of
Paniceae to transmitted phototropic excitation.
 (1) Local stimulation of the tip gives rise to two impulses, positive
and negative. The former induces a transient negative movement
(away from light); the latter causes a permanent and increasing
positive curvature towards light.
(2) Local stimulation of the growing hypocotyl gives rise to positive
curvature, subsequently neutralised by the transverse conduction of
excitation to the distal side. The absence of tropic effect in the
growing region is thus due not to lack of power of perception, but to
balanced antagonistic reactions of two opposite sides of the organ.
(3) The effects of direct and indirect stimulations are independent
of each other; hence, on simultaneous stimulations of the tip and the
growing hypocotyl, the effects of indirect stimulus are algebraically
summated with the effect of direct stimulus (neutralisation). The
indirect stimulation of the tip on the right side gives rise to two
impulses, of which the expansive positive reached the right side of the
responding region earlier, inducing convexity and movement away
from stimulus (negative curvature). This is diagrammatically shown in
Fig. 139. Had the intervening tissue been non-conducting, the slow
excitatory negative impulse would have failed to reach the responding
Fig. 139.—Diagrammatic
region, and the negative curvature induced by the positive impulse
representation of the effects of direct
and indirect stimulus on the response would prove to be the initial as well as the final effect. In the case of
of Setaria. Direct stimulation, Setaria, however, the excitatory impulse reaches the right side of the
represented by thick arrow gives rise organ after the positive impulse; the final effect is therefore an
to antagonistic concavities of induced concavity and positive curvature (movement towards
opposite sides of responding
stimulus).
hypocotyl, resulting in neutralisation.
Indirect stimulus represented by The results given above enable us to draw the following
dotted arrow gives rise to two generalisations:—
impulses, the quick positive impulse
represented by a circle, and the 1. In an organ, the tip of which is highly excitable, the balanced
slower negative impulse represented state of neutralisation, induced by direct stimulation of the responding
by crescent (concave). region, is upset in two different ways by two impulses generated in
consequence of indirect stimulation at the tip. Hence arises two types
of resultant response:—
Type A.—If the intervening tissue be semi-conducting, the positive impulse alone will reach the
growing region and induce convexity of the same side of the organ giving rise to a negative curvature.
Type B.—If the intervening tissue be conducting the transmission of the excitatory impulse will finally
give rise to a positive curvature.
Type B is exemplified by the seedling of Setaria where the transmission of excitatory impulse from
the tip upsets the neutral balance and induces the final positive curvature.
Example of type A is found in the negative phototropism of the root of Sinapis.
Negative phototropism of root of Sinapis: Experiment 140.—For investigation of the negative
phototropism of the root of Sinapis nigra I took record of its movement under unilateral action of light
by means of a Recording Microscope, devised for the purpose.[22] When the root-tip alone was
stimulated by unilateral light, the root moved away from the source of light. This was due to the
longitudinal transmission of positive impulse to the growing region at some distance from the tip. The
intervening distance between the tip and the growing region is practically non-conducting, hence the
excitatory impulse could not be conducted from the tip. After a period of rest in darkness, I next took
record of its movement under direct unilateral illumination of the growing region; the result was at first
a positive movement; but this, on account of transverse conduction of excitation under continued
stimulation, underwent a neutralisation and slight reversal. In taking a third record, in which both the
tip and growing region were simultaneously subjected to unilateral stimulation of light, I found that a
resultant responsive movement was induced which was away from light.
Thus in the root of Sinapis, the expansive effect of indirect stimulation of the tip is superposed on
that of direct stimulation of the growing region (neutral or slightly negative). The final result is thus a
movement away from light or a negative phototropic curvature.

SUMMARY.

The effect induced by stimulus of light is transmitted to a distance, in a manner precisely the same
as in other modes of stimulation.
In the Paniceae, the local unilateral stimulation of the tip of the cotyledon induces positive curvature
in the growing hypocotyl, at some distance from the tip. This is due to transmitted excitatory effect of
indirect stimulation; the earlier positive impulse induces a preliminary negative curvature, which is
reversed later by the excitatory negative impulse into positive curvature.
Contrary to generally accepted view the hypocotyl not only perceives but responds to light. The
positive curvature induced by direct stimulation is, however, neutralised by transverse conduction of
excitation.
The effects of direct and indirect stimulus are independent of each other; the final effect is
determined by their algebraical summation.
[20] Jost—Ibid—p. 468.
[21] "Response in the Living and Non-Living"—p. 17.
[22] "Plant Response"—p. 604.
Welcome to our website – the perfect destination for book lovers and
knowledge seekers. We believe that every book holds a new world,
offering opportunities for learning, discovery, and personal growth.
That’s why we are dedicated to bringing you a diverse collection of
books, ranging from classic literature and specialized publications to
self-development guides and children's books.

More than just a book-buying platform, we strive to be a bridge

connecting you with timeless cultural and intellectual values. With an
elegant, user-friendly interface and a smart search system, you can
quickly find the books that best suit your interests. Additionally,
our special promotions and home delivery services help you save time
and fully enjoy the joy of reading.

Join us on a journey of knowledge exploration, passion nurturing, and

personal growth every day!

ebookbell.com