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A new model for the archetypal plant – the perennial dicotyledonous plant (Translation of "Ein neues Modell für die Urpflanze – die mehrjährige Blütenpflanze")

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Two different models for the archetypal plant are presented. The first one is the well known model of a herbaceous, dicotyledonous plant. The second model is presented for the first time and shows a perennial, dicotyledonous plant. The choice of a model implies a particular outlook, that directs the way we look at the plant. The observer is guided. The model of the herbaceous plant takes the observer sequentially through the organs along the stem axis from the cotyledons to the carpel and induces him/her to compare those organs with each other. The new model of the perennial plant presented causes the observer additionally to compare the sprouting bud with the opened flower. The first model presents the flower that terminates the vegetative plant. The flower is related to the stem with its numerous green leaves. This is also the case for the new model, but additionally the model causes the observer to compare the flower with the sprouting bud. The leaf metamorphosis of the sprouting bud shows a clear organisation. Three regions can be distinguished: a) an area of outer sclerotic bud scales, b) an area of inner rapidly extending, distinctly colored and fragile scales and c) an area of stem leaves, just unfolding their leaf blades. This organisation is also found, as a fundamental state, in the flower. The regions correspond with a) the ring of sepals, b) the region of the petals and stamens and c) the carpel region.
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Ein neues Modell für die Urpflanze Vorabdruck // Alvaneu // 25.09.2015
Elemente der Naturwissenschaft Nr. 103, 2015. Seite 1
The following article is published in German in the journal 'Elemente der Naturwissenschaft' Nr. 103,
2015. Page 28-40. In the following paper I refer to a study on vegetative buds, that is published in
the same journal. An English version of that paper will soon be available.
Peer Schilperoord, Alvaneu 01.12.2015
DOI: 10.13140/RG.2.1.1435.1841
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A new model for the archetypal plant the perennial dicotyledonous
plant
Summary
Two different models for the archetypal plant are presented. The first one is the well known model
of a herbaceous, dicotyledonous plant. The second model is presented for the first time and shows a
perennial, dicotyledonous plant.
The choice of a model implies a particular outlook, that directs the way we look at the plant. The
observer is guided. The model of the herbaceous plant takes the observer sequentially through the
organs along the stem axis from the cotyledons to the carpel and induces him/her to compare those
organs with each other. The new model of the perennial plant presented causes the observer
additionally to compare the sprouting bud with the opened flower. The first model presents the
flower that terminates the vegetative plant. The flower is related to the stem with its numerous
green leaves. This is also the case for the new model, but additionally the model causes the observer
to compare the flower with the sprouting bud. The leaf metamorphosis of the sprouting bud shows a
clear organisation. Three regions can be distinguished: a) an area of outer sclerotic bud scales, b) an
area of inner rapidly extending, distinctly colored and fragile scales and c) an area of stem leaves, just
unfolding their leaf blades. This organisation is also found, as a fundamental state, in the flower. The
regions correspond with a) the ring of sepals, b) the region of the petals and stamens and c) the
carpel region.
Introduction
Goethe introduced the concept of the primordial or archetypal plant1 (in German: Urpflanze). He
used the concept both for a real plant that he thought he could find amidst a lot of specimens in a
garden in Italy, and for the concept of the plant, the idea, the type or the model he was constructing.
In the end it was clear to him that the archetypal plant does not exist as a real plant: "In the diaries of
my Italian journey, which are now being printed, you will, not without a smile, notice, on which
strange ways I studied the vegetative conversion; at that time I was searching for the archetypal
plant, unaware that I was looking for the idea , the concept that we need to shape the archetypal
plant".2 Goethe's innermost concern was to understand the diversity of the plants. Initially he did not
know how he could understand this diversity. The diversity confused him. He started to search for a
concept for the archetypal plant. What is the common element in all plants? He was looking for a
model and the key that belongs to the model. With both the model and the key, the two inseparable
aspects of the 'archetypal plant', it should be possible, to understand the manifold diversity of the
flowering plants.
1 in German: Urpflanze.
2 : "In den Tagebüchern meiner Italienischen Reise, an welchen jetzt gedruckt wird, werden Sie, nicht ohne
Lächeln, bemerken, auf welchen seltsamen Wegen ich der vegetativen Umwandlung nachgegangen bin; ich
suchte damals die Urpflanze, bewusstlos, daß ich die Idee, den Begriff suchte wonach wir sie uns ausbilden
könnten." (Goethe, 2006 S. 27815)
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Goethe was the founder of plant morphology with his work on the metamorphosis of plants (1790,
1817).
The term 'Urpflanze' in Goethe's sense has been forgotten. The term is used today for the very first
land plants, for the extinct ancestors of our flowering plants. Goethe uses the term as a description
of the processes common to all plants shaping their gestalt. The term 'archetypal plant' has been
replaced by terms such as type or bauplan of the angiosperms. Those terms accentuate the
schematic aspect that attaches to a 'model'. The real danger is that the key aspect is lost and that the
model becomes a kind of building set. If this is the case, it implies the use of the schemes to name
the different organs. With the concept of the 'archetypal plant' Goethe meant not only spatial, but
also temporal phenomena, the shape of the plant as the result of different processes, as the result of
different metamorphoses. The most current scientific models differ from the classical models. They
are three dimensional, computer generated with the help of mathematic formulae. These models
can simulate the growth of a specific plant. They have their justification and people who make such
models have to deal intensively with the growth of actual plants.
The crucial point in Goethe's metamorphosis teachings is to trace with attentive and inward activity
the natural growth, and after doing so to let the plant grow for the inner eye. By doing so, the inner
skills are trained, inner organs which help to understand the growth of the plant.3
Since the publication of his work many illustrations have been published in order to give an idea of
the 'archetypal plant'. The models behind these illustrations show usually one-year old dicotyledons.
The illustrations range between schematically simplified to more or less realistic. Harlan (2002)
presents an overview of 15 drawings. In this series, Wilhelm Troll's model stands out as an exception.
He presents a picture of a vegetative plant and not of a flowering plant. Troll refers erroneously to
Goethe (Schilperoord, 1997). Although the models indicate the juxtaposition of shoot and root, they
remain inaccurate in regard to the illustration of the roots, because, with one exception, the root
hairs are not shown. The majority of the models show the leaves form the cotyledon to the carpel. In
his publication Goethe's main point of view was to show the interrelationship of the leaves from the
cotyledon up to the carpel.
Models
Model of a herbaceous, dicotyledonous flowering plant
The plant (Fig. 1) shows a clear division into root and shoot. The root comprises the root axis and the
region of the root tip and root hairs. The root hairs have been drawn larger than they are in reality.
The density of the root hairs is comparable to the density of wheat root hairs. The endogenous way
of branching of the roots is exaggerated in the figure. The root branches penetrate the cortex of the
main root; the cortex being slightly domed.
3 In this connection it is important to realize that, according to Steiner (1995), thinking in childhood was once
an inward organic force of growth and that this organic force of growth transforms itself metamorphically,
emerging with the change of teeth as a soul force.
The flexibility of growth will only be fully experienced when thinking/imagining has been trained to this
flexibility.
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Figure 1. Model 1: herbaceous archetypical plant. Aquarell, 40-60 cm, painted by Franziska Fahrni-Habegger. Details in
the text. © Peer Schilperoord
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The shoot is divided into shoot axes and leaves. Leaf base, petiole and leaf blade can be
distinguished. The elongated and rounded cotyledons are followed by the first leaves. Those leaves
become stronger and stronger. One can see, that the young plant has passed through a short rosette
stage as the first four stem leaves have only short internodes, i.e. they are close together. The color
of the blades is a light green that indicates the short life span of the leaves. From the middle of the
shoot the leaves become smaller and the petiole becomes shorter. In this special case, no bracts are
formed.
We have taken garlic mustard (Alliaria petiolata), a common weed of urban areas, as the original for
the drawing of the shoot and the leaves (Fig. 2). Garlic mustard develops no bracts between the last
stem leaves and the inflorescence The plant in Fig. 1 end with a terminal flower. The flower clearly
distinguishes itself from the shoot. There is a gap between the last stem leave and the sepals.
Intermediate forms between the stem leaves and the sepals are not shown. The floral whorls are
clearly recognizable. The petals have a narrow insertion point. The number of the floral organs is
restricted. The carpel is solitary. The garlic mustard is a member of the crucifers. The gynoeceum of
the crucifers is a silique. A silique is composed of two carpels. The model shows a single, plicate
carpel. But one also regard it as a silique.
Figure 2. Part of a leaf series of a garlic mustard (Alliaria petiolata) without the cotyledons and the first stem leaves.
(Archiv Glashaus, Jochen Bockemühl)
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Model of a woody, dicotyledonous plant
As with the herbaceous plant, the one in Fig. 3 shows also a clear differentiation into root and shoot.
The lateral roots are branching. The one-year-old shoot of rhododendron served as the original. The
stem leaves are entire, the blades dark green, the cotyledons and the first stem leave are turning
yellowish.
The petiole and leaf blades of the upper stem leaves decrease in size. Unlike in the first model, the
decrease is not due to the transition to the flower but rather to the transition to the bud. It matters
not whether the bud contains a shoot or, as in Fig. 3, a flower. This metamorphosis into the bud
(from the stem leaves to the bud scales) is clearly organized. It is similar to the transition of the stem
leaves to the bracts in an annual plant.
In the axil of the next to last leaf we see a strong bud sprouting. A sprouting rhododendron shoot
served as an original for the young shoot. The metamorphosis out of the bud (from the bud scales to
the stem leaves) is totally different from the metamorphosis into the bud. Three regions are clearly
recognizable. The outer 'winter' bud scales are hard, leathery, small, full-grown and stiff; the inner
scales, extend rapidly in spring, are membranous, fall off easily, have little or no chlorophyll and can
be colored. The last region is the region of the leaves with both axis and blade. Those leaves are long
living persistent. In the figure the blades are still involuted. The transition from the inner scales to the
stem leaves often occurs abruptly.
Some small, newly formed dormant buds can be found in the axils. These buds give rise to the lateral
shoots which repeat the main shoot. The lateral buds do not originate from the apical meristem, they
arise independently at the insertion point of the leaf on the shoot axis.
The flower is in a terminal position. It was easier to draw the flower in this position and to compare it
with the sprouting shoot. The bud, which comprised the flower, is called a generative bud. From an
evolutionary point of view the terminal position is probably the more original position. The positions
of the flower and the young shoot do not have a deeper significance background. The transition from
the scales to the sepals is abrupt. Rhododendron has an inflorescence. An example of a woody plant
with terminal flowers is Camellia.
The flower is clearly organized, sepals, petals, stamens and carpels can be seen. The petals are not
grown together, they stand free like the petals of marsh marigold (Caltha palustris). The number of
the stamens corresponds with the number of the petals. The carpels are plicate, as with the follicle of
marsh marigold or like the pod of a pea. The color of the petals corresponds to the color of the inner
scales of the young shoot, as is often the case with rhododendron. One carpel is opened and shows
some ovules.
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Figure 3. Model 2: woody archetypical plant. Aquarell, ca. 40-60 cm, painted by Franziska Fahrni-Habegger. Details in the
text. © Peer Schilperoord
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Keys
Both models show the fundamental state of a dicotyledonous plant. The new model is more
generally applicable. The plants shown have reached the flowering state at the time of fertilization.
The important stage of fructification is not shown so as not to overload the pictures. At the stage of
fructification the carpel expands, the seeds develop, ripen and go into a resting phase. In the
following part the principal or key growth processes that give rise to the fully grown plants are
explained. These processes play a role in both the shaping of the vegetative plant including the
flower, and in the development of the seed.
Organic disunion
Goethe has coined the term 'organic disunion' ('organische Entzweiung') (Goethe, 1964, S. 133;
Schilperoord, 2011) Goethe writes:
"[...] A one-year-old completed plant pulled up. Ideal unity: 'When these different parts are
conceived as having arisen from an ideal body, and having developed sequentially From the very
beginning we have to consider this ideal body as being as simple as possible, and to look at it as
disunited, for without the process of disuniting of an entity, a third one cannot develop. [...]"4
The concept of organic disunion refers both to the vegetative plant and the flower. Organic disunion
already takes place at the very first development of the seed. The shoot pole and the root pole are
the result of this process. The second elementary step in the development is the division of the shoot
pole into shoot axis and stem leaves and of the root pole in root axis, root tip and, after germination,
the root hairs. Finally in the flower we can see the process of organic disunion in the development of
the pollen and the ovules.5
Iteration
An essential aspect of plant life is iteration. Leaf develops after leaf, internode follows internode,
buds are repeated. Every branch of the shoot or of the root repeats the same structures. Only at the
root tip do we find continuity; a continuing process of arising and decay.
Expansion and contraction
The process of expansion and contraction is a central motive in metamorphosis theory. After
germination, the plant has to strengthen itself it must be firmly rooted in the ground. The apical
meristem is strengthened. The leaves become taller. The next step is the decrease in size of the leaf
blade and the leaf. A new stage is announced.
Bud development and unfolding
4 The german text: "[…] Eine einjährige vollendete Pflanze ausgerauft. Ideale Einheit: Wenn diese verschiednen
Teile aus einem idealen Urkörper entsprungen und nach und nach in verschiedenen Stufen ausgebildet gedacht
werden. Diesen idealen Urkörper, mögen wir ihn in unsern Gedanken so einfach konzipieren als möglich,
müssen wir schon in seinem Innern entzweit denken, denn ohne vorher gedachte Entzweiung des einen lässt
sich kein drittes Entstehendes denken. […]"
5 The processes involving the development of the pollen and the ovules are complicated. They cannot be
shown in the picture of a flower. Morphological polarities, related with the process of organic disunion, are
clearly visible in the quality of the designs of stamen and carpel and of pollen and ovules. (Schilperoord, 1997,
2011).
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The new stage is that of the bud. The new model shows two buds opening and several dormant buds.
One bud reveals a young shoot whereas the other releases a flower. The transition of the stem leaves
into the bud scales is as, we have seen, based on a decrease in size of the blade and the petiole. until
finally only the leaf base remains in the form of the outer bud scales. The bud is a higher ranking,
superordinated organ, and is clearly differentiated and organized. The sprouting bud unveils its
differentiation, the different qualities of the tight outer winter scales, the many inner membranous
rapidly expanding scales and the strongly divergent the stem leaves.
Development of spores
The shape of the flower is related to a qualitatively new impulse. New because it is impossible to
deduce all structures of the flower from the stem leaf. The pollen sacs and the ovules are at first
sporangia, receptacles of spores. They are added to the leaf. (Schilperoord, 2011). The interaction of
the impulses of the development of the leaves and spores gives rise to stamens and carpels. The
spores develop immediately and give rise to the female gametophyte, the embryo sac and the male
gametophyte. The male gametophyte is situated in the pollen grain. These relationships need their
own illustration. We continue and look at the qualitative aspects. The extraordinary thing about
spore formation is its extreme contraction, the extreme contraction of the plant to its smallest
possible form, which is completed in the formation of the gametes. We have the smallest possible
form and the greatest formative potential. Externally, the plant disappears but is able to arise newly
after fertilization in seed formation.
Intensification
We have seen, that the development of the bud implies a heightening, a striking differentiation. This
is the moment when another impulse interacts at a very early state. The impulse to develop spores
together with the impulse to develop a bud are the basis of the spectacular diversity of flowers. In
the flower the plant is at its most individual state.
Discussion
Models describe what all plants have in common, their basic, typical characteristics. The great variety
of shapes cannot be depicted by any model. The diversity of leaf shape, of fruits and petals, has to be
elaborated in single, detailed ontogenetic studies. Ontogenesis shows the ways organs develop. The
plasticity can be experienced more intensively following the ontogenetic steps of organ development
than by comparing fully grown forms. Different shapes are the result of different ontogenetic
processes.
A typical completed leaf, out of which all other leaves can be derived, does not exist. Wilhelm Troll
and Wolfgang Hagemann have tried to find the typical angiosperm leaf shape, so to speak the
prototype for every form. That is not possible because the developmental process can take a
different direction at any moment. For a natural overview, the fully grown forms can only be related
to each other by reproducing, i.e. tracing, the ontogenetic steps (Schilperoord, 2011). An essential
part of the model is the plasticity of imagining, of thinking in processes. Otherwise the model
becomes a dead scheme. The model is tool and not an end product destination. With his model and
its key, Goethe intended to describe the essentiality of the specific plant in her liveliness.
The lively flow of organ formation can easily be seen and experienced on a plant with a richly shaped
leaf sequence from its cotyledons to its bracts and sepals. Helpful are the (rare) transitional forms,
they function like missing links to reveal the morphological relationships between the organs.
Goethe's example of hellebore species may be known. The stinking hellebore (Helleborus foetidus)
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shows transitional forms in the leaf sequences of the shoot and between stem leaves and tepals. The
Christmas rose (Helleborus niger) has no transitional forms between the last pedate stem leaf and
the first white simple tepal. The stinking hellebore is a good example for demonstrating plant
metamorphosis. Examples of transitional stages in leaf sequences in shrubs and trees are given in
Studer-Ehrensberger & Schilperoord (2015).
Geneticists like to cite Goethe, for example Enrico Coen (2000). He honored Goethe in his study
'Goethe and the ABC model of flower development'. Goethe recommends the use of malformations,
for they can reveal otherwise hidden relationships. Genetics is founded on the ability to produce
malformations. The ABC model of the geneticists describes the genetic preconditions which are
required to develop a flower in an ordinary way. How this model is related to the woody plants is to
some extent unclear and will stay unclear until the geneticists have developed a model for describing
the development of vegetative buds (Schilperoord, 2000). The ABC model has confirmed the close
relationship between petals and stamens and is in accordance with our model of the perennial
archetypal plant. Petal and stamen are the two faces of one organ. The petal emphasizes the leaf
aspect, the stamen emphasizes the spore generating aspect (Schilperoord, 2011).
Both models do not consider the great diversity of the inflorescences. With inflorescences such as
ear, panicle, umbel, or the inflorescences of the composites, plants have an enormous field of
possibilities for shaping themselves. Claßen-Bockhoff and Bull-Hereñu (2013) have developed a way
of arranging the relationships between the inflorescences by describing the ontogenetical processes
leading to the different inflorescences. They expand the model presented here by showing a a whole
spectrum of the many intermediate forms between the vegetative shoot and the single flower.
The model of the herbaceous plant differs from comparable existing by clearly emphasizing the root
hairs. The model of the woody dicotyledonous plant completes the first model by showing buds. This
is its most important feature as it shows a young shoot, developing out of a bud.
The second model permits an additional possibility for comparing plant structures with each other.
The model of the perennial plant restricts the observer to comparing the vegetative shoot with the
flower. The prevalent definition of a flower in textbooks and on the German Wikipedia website
(2015)6 is: the flower is a "unbranched short shoot with limited growth whose leaves indirectly or
directly serve sexual reproduction". The model proposed here looks at the flower as a intensified
vegetative bud. Two principal ways of metamorphosis can be recognized in the second model. The
metamorphosis from the stem leaves into the bud scales and the metamorphosis from the bud scales
into the stem leaves. The opening of the flower is comparable to the opening of a vegetative bud
(see Studer-Ehrensberger and Schilperoord, 2015). The three regions of the vegetative bud
correspond to the three regions in the flower namely the 'winter bud scales' with the sepals, the
'inner scales' with the petals and stamens and the stem leaves with the carpel region.
The intensification of the plant, is based on its metamorphosis out of the bud.
Literature
Claßen-Bockhoff, Regine; Bull-Hereñu, Kester (2013): Towards an ontogenetic understanding of
inflorescence diversity. In: Annals of Botany, S. 15231542. DOI: 10.1093/aob/mct009.
Coen, Enrico (2000): Goethe and the ABC model of flower development. In: Acad. Sci. Paris, Life
Sciences, 324 1-8.
6 not on the English site (21.09.2015)
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Elemente der Naturwissenschaft Nr. 103, 2015. Seite 11
Goethe, Johann Wolfgang von (1790): Versuch die Metamorphose der Pflanzen zu erklären. Gotha:
Ettingersche Buchhandlung.
Goethe, Johann Wolfgang von (1964): Die Schriften zur Naturwissenschaft. Bd. 10: Aufsätze,
Fragmente, Studien zur Morphologie. Bearbeitet von Dorothea Kuhn. Hermann Böhlaus
Nachfolger, Weimar.
Goethe, Johann Wolfgang von (2006): Johann Wolfgang von Goethe Leben und Werk. Digitale
Bibliothek, Berlin. Harlan, Volker (2002): Das Bild der Pflanze in Wissenschaft und Kunst. Bei
Aristoteles und Goethe, der botanischen Morphologie des 19.und 20. Jahrhunderts und bei
den Künstlern Paul Klee und Joseph Beuys. Stuttgart, Berlin: Mayer (Schriften des Karl-
Schweisfurth-Instituts für Evolutionsbiologie und Morphologie der Universität Witten-
Herdecke / Reihe B, 1).
Schilperoord-Jarke, Peer (1997): The concept of morphological polarity and its implication on the
concept of the essential organs and on the concept of the organisation of the dicotyledonous
plant. In: Acta Biotheoretica, S. 51-63.
Schilperoord-Jarke, Peer (2000): Goethes Metamorphose der Pflanzen und die moderne
Pflanzengenetik. In: Peter Heusser (Hg.): Goethes Beitrag zur Erneuerung der
Naturwissenschaften. Das Buch zur gleichnamigen Ringvorlesung an der Universität Bern zum
250. Geburtsjahr Goethes. Bern: P. Haupt, S. 131170. The English translation "Goethe's
metamorphosis and modern plant genetics" can be found on Researchgate.
Schilperoord, Peer (2011): Metamorphosen im Pflanzenreich. Stuttgart: Verl. Freies Geistesleben. S.
183. ISBN 978-3-7725-23961-5
Steiner, Rudolf (1995): The Spirit of the Waldorf School. GA297a. Anthroposophic press.
Studer-Ehrensberger, Kathrin; Schilperoord, Peer (2015): Blütenhaftes in der Metamorphose der
vegetativen Jahreszuwachseinheit der Stiel-Eiche (Quercus robur L.). In preparation:
Elemente der Naturwissenschaft, Nr. 103.
Troll, Wilhelm (1984): Gestalt und Urbild. Gesammelte Aufsätze zu Grundfragen der organischen
Morphologie. In: Böhlau Verlag, Köln, Wien.
Wikipedia (2015): https://de.wikipedia.org/wiki/Blüte 02.07.2015.
Thanks
My thanks go to Kathrin Studer-Ehrensberger for her detailed study of many buds, for the discussions
and text review and the design of the woody archetypical plant. My thanks go also to Franziska
Fahrni-Habegger for the detailed and artistic aquarelles, and last but not least to Andrea
Schilperoord ideas need time, space and talking through in order to germinate and grow.
Peer Schilperoord
Voia Gonda 1
7492 Alvaneu Dorf
schilperoord@bluewin.ch
www.urpflanze.ch
ResearchGate has not been able to resolve any citations for this publication.
Article
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Dicotyledons are polarly organised in several ways. In plant morphology polarity, a principle allowing comparison of different plant structures has until yet not been studied. A division** of the vegetative plant in shoot and root as polar structures leads to the distinction of four instead of three basic organs: leaf, shoot axis, root axis and root cap together with the root hairs. The flower is also polarly organised, its poles are formed by the carpels and the stamens. The foliage leaves are also polarly organised which is reflected by the morphological relationship of foliage leaf, stamen and carpel. The stamen uses the hypophyll*** as base of construction and the carpel uses the epiphyll**** as base of construction. Hypophyll and epiphyll are the two poles of the foliage leaf. Root and shoot, the polar entities of the vegetative plant and stamen and carpel, the polar entities of the generative plant, are morphologically correlated. Stamen and carpel can be understood as a combination of the basic organs of vegetative and generative parts of the plant. The basic organs of the generative plant are pollen grain and embryo sack with their gametophytes. The quantitative comparison of variable proportions is supplemented by a qualitative comparison of polarities. The result is, that the organisation type of the dicotyledons can yet be understood as constituted of morphological related parts. * in german Grundorgane ** in german: Gliederung *** in german: Unterblat **** in german: Oberblatt
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Zusammenfassung Das Metamorphosekonzept, das Goethe ausgearbeitet hat, bezieht sich auf das Blatt. In dieser Arbeit wird das Konzept wesentlich erweitert. Der Lebenszyklus der Pflanze wird im Sinne eines Zyklus von Metamorphosen beschrieben. Zusätzlich zu der Blattmetamorphose werden die Blattontogenese, die Embryogenese, die Trennung der Geschlechter und die verschiedenen Meristemtypen als Metamor-phosen beschrieben. Als Ergebnis der Forschungen wird anstelle der Grundor-gantheorie ein viergliedriges Modell vorgeschlagen. Weiter wird die Gestalt von Staub-und Fruchtblatt als das Ergebnis der Tätigkeit von je zwei Meristemen beschrieben. Auf die Wesensverwandtschaft zwischen Wurzel – Blattgrund – Staubblatt einerseits und Spross – Blattspreite – Fruchtblatt andererseits wird hingewiesen. Summary The concept of metamorphosis developed by Goethe is based on the leaf. In this paper the concept is substantially extended. The life-cycle of the plant is described in the sense of a cycle of metamorphoses. In addition to leaf metamorphosis, the following are also described as metamorphoses: leaf ontogenesis; embryogenesis; the separation of the sexes and the various types of meristem. As a result of these researches, instead of the fundamental organ theory, a four-component model is suggested. Furthermore, the form of the stamen and the carpel is described as the result of the activity of two meristems in each case. An indication is given regarding the relatedness of character between, on the one hand, root – leaf base – stamen and, on the other hand, shoot – leaf blade – carpel.
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Bern: P. Haupt, S. 131-170. The English translation "Goethe's metamorphosis and modern plant genetics
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Schilperoord-Jarke, Peer (2000): Goethes Metamorphose der Pflanzen und die moderne Pflanzengenetik. In: Peter Heusser (Hg.): Goethes Beitrag zur Erneuerung der Naturwissenschaften. Das Buch zur gleichnamigen Ringvorlesung an der Universität Bern zum 250. Geburtsjahr Goethes. Bern: P. Haupt, S. 131-170. The English translation "Goethe's metamorphosis and modern plant genetics" can be found on Researchgate.