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Preface

Plants are the basis of life on earth as we know it and have not only shaped, to
a large extent, diverse ecosystems but also provide food, feed, fibers, and fuel
for human subsistence. Understanding how plants evolved and develop to
fulfill their manifold functions is thus of great fundamental and applied
interest.
It will soon be 10 years since the last volume of Current Topics in
Developmental Biology focused on plant development. Since then, biology
has seen a revolution in methods for the investigation of genes, transcripts,
and proteins, allowing a wide variety of “omics” approaches. Also, the tools
to investigate gene function using forward-genetic, reverse-genetic, and
alternative methods, such as artificial microRNAs and RNA interference,
have provided deep insights into the molecular processes that regulate plant
development. The arsenal of the plant biologist has recently been extended
by the CRISPR/Cas9 system, which allows functional studies also in non-
model systems, providing new insights into developmental diversity and
plasticity. Clearly, focusing efforts on a model system, in this case Arabidopsis
thaliana, has greatly accelerated progress in understanding plant function. On
the other hand, a tunnel vision focusing on a single species is bound to miss
important aspects of development that are not found in this model system.
Certainly, plant biology has profited a lot from the powerful genetics of Zea
mays (maize), which started well over a hundred years ago, and the recent
focus on Oryza sativa (rice) as one of the most important crop plants. Over
the last years, the range of plant species used in fundamental research has
greatly expanded, providing insights into diverse developmental processes.
Unraveling the molecular control of plant development has also
provided the material to look at the evolution of gene regulatory networks,
investigating how preexisting mechanisms were co-opted for new develop-
mental programs or how novelties arose during land plant evolution. Such
evo-devo studies have provided great insights into the evolution of the
enormous diversity of plants that exists today. It is more than timely then
to provide a broad overview of plant development and evolution, as it is only
possible in a multichapter book bringing together experts in diverse field to
share their views on specific plant developmental processes and their evolu-
tion. The chapters provide a wide diversity of perspectives on both vegeta-
tive and reproductive development, summarizing the deep functional and

xvii
xviii Preface

mechanistic insights that have been gained in many fields of development.

The chapters focusing on evolutionary aspects provide an excellent view on
the directions the evo-devo field will take in the future by being able to do
functional studies in nonmodel systems that promise to provide invaluable
insights into the evolution of plant development.
I was delighted that so many of my colleagues enthusiastically accepted to
contribute to this volume of Current Topics in Developmental Biology and I am
thankful for their detailed reviews and insightful discussions on a wide range
of topics in plant development and evolution. No doubt the next decade will
see enormous progress and further deepen our understanding of plant func-
tion. Nevertheless, I am convinced that the readers will find this volume
interesting and timely and hope they will enjoy reading it.
UELI GROSSNIKLAUS
Department of Plant and Microbial Biology & Zurich-Basel Plant
Science Center, University of Zurich, Zurich, Switzerland
 CHAPTER ONE

Evolution of the plant body plan

ter Szo
Pe € ve

nyi*, Manuel Waller, Alexander Kirbis
Department of Systematic and Evolutionary Botany & Zurich-Basel Plant Science Center, University
of Zurich, Zurich, Switzerland
*Corresponding author: e-mail address: [email protected]

Contents
1. The significance of land plants 2
2. Understanding evolution of the plant body plan 2
3. Phylogenetic relationships of land plants and evolution of the land plant
body plan 3
4. Developmental patterns predating the origin of land plants 5
5. Developmental innovations of land plants 7
5.1 Alternation of haploid (gametophyte) and diploid (sporophyte)
generations 7
5.2 Evolution of three-dimensional growth in the haploid and diploid phases 10
5.3 Origin of spores, sporangia, and sporopollenin in land plants 12
5.4 Origin of unbranched sporophyte forms 12
5.5 Evolution of bifurcating axes 14
5.6 Evolution of indeterminacy 15
5.7 Evolution of meristems 16
5.8 Origin of leaves 17
5.9 Evolution of rooting systems 20
5.10 Roots 22
6. Conclusions and perspectives 24
Acknowledgments 25
References 25

Abstract
Land plants evolved about 470 million years ago or even earlier, in a biological crust-
dominated terrestrial flora. The origin of land plants was probably one of the most sig-
nificant events in Earth’s history, which ultimately contributed to the greening of the
terrestrial environment and opened up the way for the diversification of both plant
and non-plant lineages. Fossil and phylogenetic evidence suggest that land plants have
evolved from fresh-water charophycean algae, which were physiologically, genetically,
and developmentally potentiated to make the transition to land. Since all land plants
have biphasic life cycles, in contrast to the haplontic life cycle of Charophytes, the
evolution of land plants was linked to the origin of a multicellular sporophytic phase.
Land plants have evolved complex body plans in a way that overall complexity

Current Topics in Developmental Biology, Volume 131 # 2019 Elsevier Inc. 1

ISSN 0070-2153 All rights reserved.
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2 P
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increased toward the tip of the land plant tree of life. Early forms were unbranched, with
terminal sporangia and simple rhizoid rooting structures but without vasculature and
leaves. Later on, branched forms with lateral sporangia appeared and paved the route
for the evolution for indeterminacy. Finally, leaves and roots evolved to enable efficient
nutrient transport to support a large plant body. The fossil record also suggests that
almost all plant organs, such as leaves and roots, evolved multiple times independently
over the course of land plant evolution. In this review, we summarize the current knowl-
edge on the evolution of the land plant body plan by combining evidence of the fossil
record, phylogenetics, and developmental biology.

1. The significance of land plants

Land plants evolved about 470 million years ago, or potentially even
earlier, in the Cambrian (515 Ma) to Early Ordovician (473 Ma) when the ter-
restrial flora was dominated by a crust-forming assemblage of algae lichens,
bacteria, cyanobacteria, and fungi (Berbee, James, & Strullu-Derrien, 2017;
Morris et al., 2018; Salamon et al., 2018). The origin of land plants was
one of the most significant events in the history of the globe, which paved
the road for diversification of non-plant lineages in the terrestrial environment
(Bateman et al., 1998; Berner, 1997). Among others, this was enabled by
an increasing oxygen concentration produced through photosynthesis, ini-
tially by cyanobacteria and then eukaryotic algae and land plants (Kenrick,
Wellman, Schneider, & Edgecombe, 2012). The resulting ozone layer
reduced the amount of UV irradiation, which was probably necessary for
plants to leave the water (Rozema et al., 2002; Rozema, Blokker, Mayoral
Fuertes, & Broekman, 2009). Furthermore, land plants have also significantly
contributed to the weathering of rocks and the formation of soil by preventing
transportation of sediments by wind and water (Lenton, Crouch, Johnson,
Pires, & Dolan, 2012; McMahon & Davies, 2018; Perkins, 2018). Altogether,
plants have tremendously contributed to the greening of the planet and to the
rapid increase of terrestrial biodiversity.

2. Understanding evolution of the plant body plan

Three major types of information can be utilized to investigate the
main trends in the evolution of the land plant body plan, and to uncover
the underlying genetic mechanisms. Phylogenetic analyses of extant land
plants can provide information on the ancestor-descendant relationships
of major lineages, and thus a rough timeline of their evolutionary origin.
Evolution of the plant body plan 3

Nevertheless, available phylogenetic evidence is mostly based on nucleotide

data and includes only extant plants, providing little information about the
combined evolutionary history of extant and fossil forms.
Another line of evidence is provided by the fossil data. Unfortunately,
fossil record of the first land plants is sparse, and their interpretation can
be ambiguous (Boyce & Kevin Boyce, 2010; Kenrick, 2018; Tomescu,
2009). Ambiguity in the assessment of morphological characters and their
potential homology with body plans of extant land plants make the evolu-
tionary placement of fossils questionable and combined analysis of data on
fossils and extant land plants is challenging (Bateman, 1996; Puttick et al.,
2018; Salamon et al., 2018; Stewart & Rothwell, 1993; Seward, 2011).
Despite this, careful comparative morphological and evolutionary analyses
of the fossil record and extant land plants gave rise to major theories about
the evolutionary origin of main modular units of the land plant body plan,
providing testable hypotheses (Chomicki, Coiro, & Renner, 2017).
Comparative analysis of fossils and phylogenetic relationships of extant
land plants is necessary to reveal evolutionary trends and formulate testable
hypotheses, but do not allow experimental validation. Comparative devel-
opmental biology and genetics/genomics of extant representatives of land
plants can be used to experimentally test hypotheses on the developmental
and genetic mechanisms underlying the evolutionary trends in land plant
body plan evolution (Boyce & Kevin Boyce, 2010; Rothwell, Wyatt, &
Tomescu, 2014; Tomescu, Wyatt, Hasebe, & Rothwell, 2014).
In this review, we provide insights into the recent advancement surround-
ing the evolution of the land plant body plan by evaluating evidence from
phylogenetics/phylogenomics, the fossil record, comparative genomics, and
the developmental biology of plants. We do so by summarizing recent
evidence in these scientific fields and their implications for each major inno-
vation in the evolution of the land plant body plan (see Fig. 1).

3. Phylogenetic relationships of land plants

and evolution of the land plant body plan
Phylogenetic analysis of extant members of green plants unambigu-
ously suggests that the lineage of land plants (embryophytes) together with
several lineages of streptophytic algae (charophycean algae) forms the mono-
phyletic group of streptophytes (Becker & Marin, 2009; Gitzendanner,
Soltis, Wong, Ruhfel, & Soltis, 2018; Liu, Cox, Wang, & Goffinet,
2014; Wickett et al., 2014; Wodniok et al., 2011). The streptophytes are
4 P
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enyi et al.

Fig. 1 Hypothetical phylogenetic relationship of extant and fossil land plants and Char-
ophytes. The left bar shows the timescale of evolution. Names of extant groups are in
black while extinct taxa/lineages are shown in gray. Dashed horizontal lines represent
ambiguous phylogenetic relationships. Dotted vertical lines indicate the uncertainty in
the timing of the origin of embryophytes and lineages of bryophytes. Current data using
phylogenetic dating suggests that the deep splits in the tree may be considerably older
than depicted here (Morris et al., 2018). Pictograms show the architectural features of
major plant groups. Open elliptic structures refer to sporangia while gray filled structures
depict leaves. Redrawn, extended and modified from Kenrick, P., & Crane, P. (1997). The origin
and early diversification of land plants: A Cladistic Study. Science, 389(4), 33–39.

further embedded into the large clade of the green lineage (green plants
including green algae), the Viridiplantae. Three lineages of the paraphyletic
grade of streptophyte algae, the Charophyceae, Coleochaetophyceae, and
the Zygnematophyceae, are the closest relatives of land plants and share
the presence of phragmoplast with land plants, a structure central to the for-
mation of a new cell wall after cell division. Of the streptophyte algal line-
ages, the Zygnematophyceae appear to be the closest relative of land plants
(Delwiche & Cooper, 2015; Timme, Bachvaroff, & Delwiche, 2012;
Wickett et al., 2014; Wodniok et al., 2011). Extant Zygnematophyceae
consist of unicellular and filamentous algae, occupying freshwater and ter-
restrial habitats, whose developmental features are very difficult to compare
with the complexity of land plant body plans (Delwiche & Cooper, 2015).
Evolution of the plant body plan 5

Therefore, having the Zygnematophyceae as sister to land plants makes recon-

struction of the character states of the shared common algal ancestor with land
plants particularly challenging (de Vries & Archibald, 2018). Altogether, the
sister relationship of Zygnematophyceae with land plants provides very little
information on the potential order and extent of evolutionary transformations
that led to the evolution of the complex body plans of early land plants.
Phylogenetic analysis of extant land plants (embryophytes) implies that
land plants consist of the clade of vascular plants, which include the monophy-
letic groups of lycophytes, monilophytes, gymnosperms and angiosperms,
and a group of three lineages, the mosses, liverworts, and hornworts, collec-
tively referred to as bryophytes (Cox, Li, Foster, Martin Embley, & Civáň,
2014; Gitzendanner et al., 2018; Liu et al., 2014; Wickett et al., 2014;
Wodniok et al., 2011). The phylogenetic inter-relationships of the bryophyte
lineages and their relationship to vascular plants are still highly debated. For
instance, earlier analyses, mainly based on molecular data, recovered all pos-
sible topologies between the three lineages of bryophytes and the rest of land
plants. Either mosses, hornworts, or liverworts were reconstructed as sister to
the rest of the land plants, and the three lineages of bryophytes were mainly
resolved as a paraphyletic grade with some exceptions (Bremer, Humphries,
Mishler, & Churchill, 1987; Chang & Graham, 2011; Cox et al., 2014; Finet,
Timme, Delwiche, & Marl
etaz, 2010; Fiz-Palacios, Schneider, Heinrichs, &
Savolainen, 2011; Karol, 2001; Laurin-Lemay, Brinkmann, & Philippe, 2012;
Mishler & Churchill, 1984; Qiu et al., 2006; Wodniok et al., 2011; Zhong,
Liu, Yan, & Penny, 2013). Conversely, recent phylogenomic analyses impli-
cate that mosses and liverworts are very likely monophyletic (Gitzendanner
et al., 2018; Liu et al., 2014; Morris et al., 2018; Puttick et al., 2018;
Wickett et al., 2014; Wodniok et al., 2011). Furthermore, some analyses give
strong support to the monophyly of bryophytes revealing the deepest split
between hornworts and a clade consisting of the liverworts and mosses
(Cox et al., 2014; Gitzendanner et al., 2018; Liu et al., 2014; Wickett
et al., 2014; Wodniok et al., 2011; Zhong et al., 2013).

4. Developmental patterns predating the origin

of land plants
Recent evidence suggests that the genetic basis for cellular processes that
allowed plants to colonize and successfully cope with the terrestrial environ-
ment were already present in the charophycean algae and, therefore, their
evolution predated the origin of land plants (de Vries & Archibald, 2018;
6 P
eter Sz€
ov
enyi et al.

de Vries, Curtis, Gould, & Archibald, 2018; Harholt, Moestrup, & Ulvskov,
2016; Selosse, Strullu-Derrien, Martin, Kamoun, & Kenrick, 2015). Similarly,
it was found that most transcription factor families, key regulators of
developmental processes thought to be specific to land plants, were already
present in the charophycean algae (Catarino, Hetherington, Emms,
Kelly, & Dolan, 2016; Wilhelmsson, M€ uhlich, Ullrich, & Rensing, 2017).
This suggests that some developmental innovations relevant to land plant evo-
lution may have been already present in the charophycean algae.
Charophycean algae are highly diverse in their development. For
instance, some are unicellular, but others have attained multicellularity with
thallose or filamentous forms (Delwiche & Cooper, 2015; Domozych,
Popper, & Sørensen, 2016). They may or may not show apical growth,
with some groups exhibiting complex apical growth and branching. There-
fore, developmental patterns such as apical growth, branching, and
multicellularity were present already in the charophycean algae. It is assumed
that some of these developmental patterns, and likely the corresponding
mechanisms, were retained through the evolution of early land plants. For
instance, the thallose gametophytes of liverworts resemble those of the char-
ophytes, and the underlying developmental mechanisms may have been pre-
sent and retained from their common ancestor (Ligrone, Duckett, &
Renzaglia, 2012a; Renzaglia, Duff, Nickrent, & Garbary, 2000). Similarly,
the presence of both unicellular and multicellular forms in charophytes sug-
gests that developmental processes for multicellularity might have been pre-
sent in the common ancestor of charophytes and land plants and retained.
Intriguingly, some experiments suggest that the multicellular-unicellular
transition can be achieved in moss protonema by targeting genes involved
in protein prenylation. Such mutants have unicellular cells undergoing unpo-
larized divisions and resemble those of unicellular algal cells (Antimisiaris &
Running, 2014; Thole, Perroud, Quatrano, & Running, 2014). Further-
more, recent experiments with the alga Chlamydomonas reinhardtii suggest that
multicellularity can be relatively easily attained under certain selection regimes
or by altering the regulation of cell cycle genes (Boyd, Rosenzweig, & Herron,
2018; Hanschen et al., 2016; Herron, 2016; Niklas, 2014; Olson & Nedelcu,
2016; Ratcliff et al., 2013). The observation that the basic genetic tool kit of
land plants is present in the genome of the charophyte Klebsormidium flaccidum,
and that most key transcription factor families are present in various char-
ophycean algae, further suggest that a set of developmental mechanisms and
their genetic regulators were already present in the common ancestor of
charophytes and land plants (Hori et al., 2014; Wilhelmsson et al., 2017).
Evolution of the plant body plan 7

Altogether, this implies that the common ancestor of land plants and char-
ophytes was genetically and likely developmentally prepared to achieve the
complex body plan of land plants through relatively few evolutionary steps.
Nevertheless, except some studies on MADS box genes (Tanabe et al.,
2005), information on the genetic regulation of specific developmental pro-
cesses of charophytes is lacking, and future research will provide information
about the potential homology of basic developmental mechanisms in land
plants and charophytes.

5. Developmental innovations of land plants

Transition to the terrestrial environment brought some radical changes
to the basic body plan of plants. The appearance of biphasic life cycles, spo-
ropollenin and sporangia, three-dimensional growth and simple unbranched
sporophytes appeared very early in land plant evolution.

5.1 Alternation of haploid (gametophyte) and diploid

(sporophyte) generations
All charophycean algae had unicellular ancestral forms and a life cycle in which
the haploid phase dominates, and the syngamy of gametes is immediately
followed by meiosis of the zygote (Bowman, Sakakibara, Furumizu, &
Dierschke, 2016; Niklas & Kutschera, 2009; Qiu, Yin-Long, Taylor, &
McManus, 2012). Such life cycles are haplontic because all mitotic divisions
occur in the haploid phase, and the diploid phase is only represented by a sin-
gle cell, the zygote. The body plan of land plants radically broke this pattern
by evolving multicellular alternating haploid (gametophytic) and diploid
(sporophytic) phases. Relative dominance of the two phases has also changed
during the course of land plant evolution, in such a way that the haploid phase
dominated early in evolution, followed by the elaboration of the diploid and
reduction of the haploid phase later in evolution (Bowman et al., 2016;
Niklas & Kutschera, 2009; Qiu et al., 2012). Therefore, the origin of land
plants is linked to the evolution of the multicellular sporophytic phase.
Current phylogenetic evidence, consistently resolving charophycean algae
with a purely haplontic life cycle as sister to all land plants, gives overwhelming
support to this assertion. This finding is consistent with the antithetic theory
proposing that land plants evolved from an algal ancestor with a haplontic life
cycle with zygotic meiosis. In parallel, phylogenetic evidence rejects the
homologous theory of Bower, assuming an algal ancestor with an isomorphic
alternation of haploid and diploid generations. Altogether, current evidence
Visit https://ebooksecure.com
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8 P
eter Sz€
ov
enyi et al.

implies that the multicellular sporophyte was an evolutionary innovation of

land plants and originated by the intercalation into the ancestral haplontic life
cycle of mitotic divisions in the zygote prior to meiosis (Bower, 1890;
Haig, 2008).
Therefore, the major difference between the haplontic life cycles of
charophytes and the biphasic life cycles of land plants is whether the zygote
proceeds to meiosis without mitotic divisions or it proliferates mitotically
prior to meiosis. Intensive research on this topic over the last years suggests
that part of the genetic toolkit responsible for this developmental switch seems
to be deeply rooted in the history of the green lineage (Bowman et al., 2016).
Theory further suggests that the evolution of this toolkit may date back to the
origin of green algal mating types, which likely evolved to impose a stringent
control on the timing of developmental switches (Wang & Dohlman, 2005).
The molecular toolkit controlling these critical developmental aspects of the
biphasic life cycle of land plants seems to have evolved by co-opting
the genetic network controlling meiosis in the zygote of simple unicellular
algae, such as C. reinhardtii (Bowman et al., 2016; Lee, Lin, Joo, &
Goodenough, 2008). In C. reinhardtii, heterodimerization of the BEL1-LIKE
(BELL) homeodomain transcription factor GSP1 with the KNOTTED1-
LIKE HOMEOBOX (KNOX) family transcription factor GSM1 is necessary
to initiate zygotic gene expression and meiosis (Bowman et al., 2016; Lee
et al., 2008). Homologs of both BELL and KNOX gene families are present
in all land plants and they do heterodimerize (Floyd & Bowman, 2007;
Frangedakis, Saint-Marcoux, Moody, Rabbinowitsch, & Langdale, 2017;
Hay & Tsiantis, 2010; Horst et al., 2016). KNOX and BELL function is best
studied in the moss Physcomitrella patens, in which a single BELL gene,
PpBELL1, is sufficient and necessary to induce the sporophytic program.
It is also known that KNOX genes went through a duplication prior to
the origin of land plants, giving rise to Class 1 and Class 2 KNOX genes
(Floyd & Bowman, 2007; Frangedakis et al., 2017; Furumizu, Alvarez,
Sakakibara, & Bowman, 2015; Hay & Tsiantis, 2010). The Class 1 KNOX
gene of P. patens, MKN2, is necessary for regulating sporophyte development
in the moss, while the Class II KNOX genes appear to suppress the gameto-
phytic program in the sporophytic phase (Sakakibara et al., 2013; Sakakibara,
Nishiyama, Deguchi, & Hasebe, 2008; Sano et al., 2005). It is not yet
clear with which KNOX protein PpBELL1 heterodimerizes with (Horst
et al., 2016). Conversely, components of the Polycomb Repressive
Complex 2 (PRC2), such as the homologs of the Arabidopsis thaliana
proteins FERTILIZATION-INDEPENDENT ENDOSPERM (PpFIE)
Evolution of the plant body plan 9

and CURLY LEAF (PpCLF), suppress the sporophytic program in the game-
tophytic phase (Mosquna et al., 2009; Okano et al., 2009; Pereman et al.,
2016). Furthermore, transcriptomic data suggest early activation of meiosis-
related genes in the apical cells of the sporophyte in the moss P. patens
(Frank & Scanlon, 2015a, 2015b). Nevertheless, the link between the gene
networks regulating the haploid-diploid switch and that of sporangial
development is unknown. These findings suggest that an ancient regulatory
network, whose evolution coincided with the origin of mating types in green
algae, is deeply conserved across the green lineage. The putative ancestral
function of this network was to enable a stringent control on the developmen-
tal switch between haploid and diploid programs (Bowman et al., 2016).
This network was then used, and its regulatory role was extended, to govern
various aspects of sporophytic development in land plants.
Although the core regulatory network controlling initiation of the sporo-
phyte and gametophyte developmental programs is known, there is very little
information available about the genetic program enabling multicellularity and
three-dimensional growth in the sporophytic phase. It is possible that these
developmental mechanisms were partly recruited from the gametophytic
phase (Frank & Scanlon, 2015a, 2015b; Szovenyi, Rensing, Lang, Wray, &
Shaw, 2010). Alternatively, they could have evolved de novo in land plants.
It is known that zygotes of double mutants disrupting the FLORICAULA/
LEAFY (FLO/LFY) homologs PpLFY1 and PpLFY2 arrest and are unable
to divide mitotically (Tanahashi, Sumikawa, Kato, & Hasebe, 2005). Further-
more, P. patens zygotes lacking activity of the two WUSCHEL-RELATED
HOMEOBOX 13-LIKE (PpWOX13LA/B) genes are unable to elongate and
initiate the apical cell of the embryo (Sakakibara et al., 2014). Transcriptomic
evidence from P. patens suggests that multicellularity and three-dimensional
patterning may have been, at least partially, recruited from gametophytic pro-
grams (Frank et al., 2015; Frank & Scanlon, 2015b; Whitewoods et al., 2018).
Nevertheless, many aspects of the evolution of the sporophytic phase are still
unclear, and the interconnection of reproductive and proliferative programs is
unknown.
Although, multiple lines of evidence support the origin of land plants
from a charophycean ancestor, there is much ambiguity surrounding the
evolutionary relationship of land plant lineages. The debate about the phy-
logenetic relationship of bryophyte lineages and their relationship with
vascular plants has significant impact on how land plant body plan changes
are interpreted. This includes the evolution of the biphasic life cycle and the
evolution of other morphological traits, such as stomata, vascular tissues, etc.
10 P
eter Sz€
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enyi et al.

For instance, studies suggesting a paraphyletic relationship of the three

bryophyte lineages (Chang & Graham, 2011; Qiu et al., 2006, 2012) are com-
patible with a haploid-dominant ancestral land plant life cycle, which was
retained in the paraphyletic grade of bryophytes but was followed by the
dominance of the diploid phase in vascular plants (Bowman et al., 2016;
Haig, 2008; Niklas & Kutschera, 2009; Qiu et al., 2012; Tomescu et al.,
2014). In contrast, a monophyletic bryophyte clade may suggest that the
haploid-dominant life cycle could have been a unique innovation of the bryo-
phyte clade (Cox et al., 2014; Gitzendanner et al., 2018; Puttick et al., 2018;
Wickett et al., 2014). According to this hypothesis, life cycle of the ancestral
land plant could have been haploid-dominant, diploid-dominant, or equally
dominant with close to isomorphic haploid and diploid phases. Furthermore,
it is possible that the common ancestor of all land plants may have been more
trachaeophyte-like, which may explain the origin of conducting tissues in
mosses. The earliest fossil remains of land plants are currently interpreted as
being stem trachaeophytes (Kenrick, 1994, 2018; Kenrick & Crane, 1997;
Taylor, Kerp, & Hass, 2005). Nevertheless, it is possible that some of these
fossils represent the common ancestor from which the monophyletic group
of bryophytes evolved (Boyce & Kevin Boyce, 2010; Kenrick, 2018;
Puttick et al., 2018). Therefore, there is considerable uncertainty surrounding
the evolutionary origin of land plants, which makes reconstruction polarity of
key characters in land plants challenging.

5.2 Evolution of three-dimensional growth in the haploid

and diploid phases
Most charophycean algae grow in a planar form along a two-dimensional axis,
while land plants evolved three-dimensional growth, enabled by the presence
of a continuously rotating division plane in the stem cells (Delwiche &
Cooper, 2015; Domozych et al., 2016; Langdale, 2008). The ability of flexibly
changing division planes is missing from the algal relatives of land plants. The
genetic mechanisms underlying this evolutionary transition can only be stud-
ied in land plants with a life cycle including both two- and three-dimensional
growth patterns.
Importantly, some mosses exhibit a filamentous juvenile life cycle phase
(protonemata) of the gametophyte, which is maintained by two-dimensional
growth as seen in charophycean algae. After this stage, a change to three-
dimensional growth may occur, giving rise to leafy shoots. This system is
suitable to investigate the genetic mechanisms enabling the transition from
Evolution of the plant body plan 11

two- to three-dimensional growth. Nevertheless, this approach assumes that

the ontogeny of the moss P. patens recapitulates the evolutionary trajectory
that occurred between algae and land plants (Harrison, Roeder,
Meyerowitz, & Langdale, 2009). There are multiple genetic factors known
to affect initiation of the three-dimensional growth pattern in P. patens.
The NO GAMETOPHORES1 (PpNOG1) gene regulates the transition to
three-dimensional growth by inducing the degradation of proteins that likely
repress the P. patens APB transcription factors, belonging to the AP2 class tran-
scription factors homologous to A. thaliana AINTEGUMENTA, PLETH-
ORA, and BABY BOOM (APB). The PpAPB transcription factors are
necessary and sufficient to initiate three-dimensional buds (Aoyama et al.,
2012; Moody, Kelly, Rabbinowitsch, & Langdale, 2018) The PpNOG1 gene
is likely also necessary in directing division planes in the apical cell, together
with the DEFECTIVE KERNEL1 homolog PpDEK1, to achieve proper
three-dimensional growth (Demko et al., 2014; Olsen, Perroud,
Johansen, & Demko, 2015; Perroud et al., 2014). PpNOG1 and related genes
occur only in land plants; therefore, it may be one of the key factors that reg-
ulate this innovation (Moody et al., 2018). Given that PpDEK1 and AP2 class
transcription factors regulate similar processes in flowering plants and the
moss, it is possible that this ancient network was already present in the com-
mon ancestor of land plants, and that its evolution coincided with the evolu-
tion of three-dimensional growth. Finally, homologs of the CLAVATA
(CLV) pathway, crucial for meristem maintenance in the flowering plant
shoot apical meristem, were recently shown to be critical to the correct ori-
entation of division planes in the transition from two- to three-dimensional
growth in P. patens (Whitewoods et al., 2018). The role of the CLV pathway
in orienting division planes seems to be conserved between A. thaliana and
P. patens and may represent the ancient function of this pathway. It is thus
possible that the CLV pathway was recruited to regulate the proper orienta-
tion of division planes in two independent contexts: in the apical cell of the
moss gametophyte and in the shoot apical meristem of flowering plants. Phe-
notypic effects of PpDEK1, PpNOG1 and the moss homologs of the CLV
pathway are overlapping, suggesting that they are likely members of a gene
network governing the proper development of the gametophyte apical cell
in P. patens. In line with these findings, transcriptomic evidence also suggests
that the genetic mechanisms involved in the evolution of three-dimensional
growth in gametophytes may have been partially recruited to support three-
dimensional patterning in the sporophytic phase (Frank & Scanlon, 2015a).
12 P
eter Sz€
ov
enyi et al.

5.3 Origin of spores, sporangia, and sporopollenin

in land plants
All land plant spores are covered by sporopollenin, whereas algae only have
heavy-walled zygotes that germinate via meiosis. Therefore, sporopollenin
coated spores are an innovation specific to land plants. This assertion is in line
with the finding that the earliest fossils with land plant affinities are the
so-called cryptospores (470 million years old), whereas sporangial fragments
are younger (450 million years old) (Edwards, Morris, Richardson, &
Kenrick, 2014; Steemans et al., 2009). Cryptospores occur either in monads,
diads, or in tetrads (see chapter “Evolution and co-option of developmental
regulatory networks in early land plants” by Bowman et al., this issue). Their
wall structure resembles that of some liverwort spores, such as the diads and
tetrads of the genera Haplomitrium and Sphaerocarpos, respectively, but their
affinity to extant bryophyte species is highly debated (Edwards, Richardson,
Axe, & Davies, 2012; Gensel, 2008; Renzaglia et al., 2015). It is thought that
sporopollenin-covered spores evolved earlier than sporophytes, likely via
modification of the timing of meiosis and the deposition of sporopollenin,
which significantly affected spore viability (Brown & Lemmon, 2011).
Genetic pathways and their components are partially conserved across land
plants, but conservation of the genetic components underlying the develop-
mental process and its evolutionary trajectory is not known (Harrison,
Alvey, & Henderson, 2010).

5.4 Origin of unbranched sporophyte forms

The earliest non-spore fossils assigned to land plants are believed to resemble
present day vascular plants with bifurcating axes (Bowman, 2013; Boyce &
Kevin Boyce, 2010; Kenrick, 2018; Kevin Boyce, 2005; Gensel, 2008;
Langdale, 2008). Nevertheless, phylogeny and the earliest fossils of stem
group polysporangiophytes suggest that early sporophytes may have been
unbranched, with terminal sporangia such as in extant mosses. Furthermore,
the earliest sporophytes are believed to be obligate matrotrophic (nutrition-
ally supported by the gametophyte), with a well-developed photosynthetic
gametophyte phase (Gensel, 2008; Kenrick, 2018; Qiu et al., 2012; Remy,
Gensel, & Hass, 1993). Therefore, it is hypothesized that fossils of the earliest
multicellular unbranched sporophyte forms are missing from the fossil
record, as all known fossils show some bifurcation. Nevertheless, their small
size and the lack of vascularization correspond to the organizational level
of bryophyte sporophytes (Boyce & Kevin Boyce, 2010; Gensel, 2008;
Evolution of the plant body plan 13

Gerrienne et al., 2006; Remy et al., 1993; Taylor et al., 2005). It is assumed
that these unbranched forms are retained in extant bryophytes, especially in
the lineages of mosses and hornworts, which can be used to gain information
about their developmental biology and evolution. Unfortunately, develop-
ment of the sporophyte in the three lineages of bryophytes is strikingly dif-
ferent and, therefore, it is difficult to establish which features are ancestral to
the group (Ligrone et al., 2012a; Ligrone, Duckett, & Renzaglia, 2012b)
(Fig. 2).
Importantly, elongation of the liverwort sporophyte is almost exclusively
due to cell expansion, and there is no apical or localized meristem activity with
divisions occurring all over the sporophyte (Ligrone et al., 2012a; Renzaglia
et al., 2000). In contrast, sporophyte growth is dominated by cell divisions in
both hornworts and mosses, involving the activity of well-localized intercalary
meristems and/or apical cells (Ligrone et al., 2012a; Villarreal & Renzaglia,
2015). Hornwort and moss sporophytes share the presence of a so-called mul-
ticellular proliferative zone within the sporophyte (Ligrone et al., 2012b).
This proliferative zone produces cells upward, giving rise to the full body
of the sporophyte in hornworts, whereas it produces cells downward in

Fig. 2 Schematic representation of sporophyte development in the three lineages of

bryophytes. The bottom row shows the three cell layers (basal, middle and upper) of
the early embryo. The upper row depicts the developing sporophytes. Actively dividing
cells are shown in gray and arrows refer to the direction of cell production. Inactive but
preformed regions are shown in black. Randomly arranged arrows in the liverwort
embryo symbolize cell divisions occurring across the whole embryo. Gray arrow heads
in the developing sporophyte of liverworts depict cell elongation and a lack of cell divi-
sions leading to the elongation of the seta.
14 P
eter Sz€
ov
enyi et al.

mosses, giving rise to the stalk (seta) of the sporophyte (French & Paolillo,
1975b; Villarreal & Renzaglia, 2015). Developmental homology of these
structures and their relation to the probably ancient uniaxial structures is
unexplored. Nevertheless, historical experiments of moss sporophytes suggest
that cytokinin may be involved in regulating activity of the intercalary meri-
stem. Retarded growth of the sporophyte after decapitation can be alleviated by
exogenous cytokinin application, which may represent a conserved mechanism
with the angiosperms shoot apical meristem (SAM) (French & Paolillo, 1975a,
1975b; Ligrone et al., 2012b; Snipes et al., 2018; Truskina & Vernoux, 2018).
It is not yet clear, how developmental mechanisms of unbranched forms
are linked to the origin of branched shoots of early vascular plants. There are
two competing hypotheses explaining the evolutionary origin of early vascular
shoot systems, the interpolation/sterilization and apical growth hypotheses
(Bowman, 2013; Ligrone et al., 2012b; Tomescu et al., 2014). The apical
growth hypothesis assumes that early shoots evolved from the extended mer-
istematic activity of the transient apical cell of a moss-type embryo. In contrast,
the interpolation/sterilization hypothesis suggests that the common ancestor
of bryophyte and vascular plant sporophytes had apical reproductive activity,
which went through a sterilization process to produce the axis of the shoot.
Further comparative study on the unbranched sporophytes of bryophytes and
that of vascular plants may clarify whether any of these hypotheses can be
supported by experimental data.

5.5 Evolution of bifurcating axes

The earliest vascular plant remains had independent gametophyte and spo-
rophyte generations, with bifurcating sporophyte axes that terminated in
sporangia. This is in contrast to the unbranched and matrotroph sporophytes
of extant mosses (Gensel, 2008; Kenrick, 2018; Remy et al., 1993; Taylor
et al., 2005; Tomescu et al., 2014). Indeterminate sporophytes appear only
much later in the fossil record, implying that evolution of bifurcation pre-
dates that of indeterminacy. Information on how branching evolved is
provided by observations on fern and bryophyte sporophytes. In P. patens
plants, deletion either of the TEOSINTE BRANCHED1/CYCLOIDEA/
PROLIFERATING CELL FACTOR1 (TCP) transcription factor class
gene PpTCP5, the TERMINAL EAR1-LIKE (TEL) gene PpTEL, or
PINFORMED1 (PIN1) homolog PpPINB gene will increase the proportion
of bifurcating sporophytes compared to the wild type (Bennett et al., 2014;
Ortiz-Ramı́rez et al., 2016; Vivancos et al., 2012). PpFLO/LFY mutants have
Evolution of the plant body plan 15

a similar phenotype (Tanahashi et al., 2005). At present, it is unclear when

exactly branching takes place and whether it occurs at the zygote or at the
sporophyte-seta stage. It is also unclear whether the effect of these genes is
achieved via similar developmental mechanisms. Observations on fern sporo-
phytes show that branching is achieved by segregation and amplification of
stem cells at the shoot apex (Harrison, Rezvani, & Langdale, 2007). Trans-
criptomic evidence suggests that genetic components necessary for this process
are present in fern genomes and shoot apices, but the genetic mechanisms are
still unclear (Evkaikina et al., 2017; Frank et al., 2015; Harrison, 2015). In
contrast, gametophytic branching in mosses is regulated by the co-option
of ancient hormonal effects involved in sporophyte branching in angiosperms,
such as auxin, cytokinins, and strigolactons (Coudert, Bell, Edelin, &
Harrison, 2017; Coudert et al., 2015). Polar auxin transport is essential for
branching in flowering plants but in the moss bi-directional transport is
required for normal branching (Coudert, 2017; Harrison, 2017).

5.6 Evolution of indeterminacy

The fossil record suggests that all indeterminate sporophyte axes have later-
ally arranged sporangia (Boyce & Kevin Boyce, 2010; Tomescu et al., 2014).
Therefore, the evolution of indeterminacy of sporophytic axes and the
lateral displacement of sporangia are linked, indicating spatially and tempo-
rally separated activity of reproductive and vegetative functions (Kenrick,
2018). Information on the molecular mechanisms underlying indeterminacy
is exclusively coming from investigations on the determinant sporophyte of
P. patens. It was shown that two components of the moss PRC2, encoded by
PpFIE and PpCLF, are necessary to repress the meristematic activity of the
sporophyte apical cell (Kenrick, 2018; Mosquna et al., 2009; Pereman et al.,
2016). In mutants disrupting PRC2, the sporophytes produced branched
structures by continuous proliferation. Class I KNOX genes are known
to be responsible for the maintenance of the meristematic activity of the spo-
rophyte apical cell (Sakakibara et al., 2008). That is, the interaction of PRC2
and Class I KNOX gene activity is key in the regulation of determinant and
indeterminant growth. Class II KNOX genes are also expressed in and nec-
essary for the development of the sporophyte by repressing the gameto-
phytic program (Sakakibara et al., 2013). These observations imply that
function of the PRC2 in repressing pluripotent sporophytic cells is con-
served across land plants. Whether the antagonistic effect of Class I and II
KNOX genes seen in flowering plants is also conserved in the development
16 P
eter Sz€
ov
enyi et al.

of the sporophyte of mosses is unknown (Furumizu et al., 2015). Neverthe-

less, it is possible that evolution of indeterminacy is partially achieved by the
proper coregulation of Class I and Class II KNOX activities, which may
connect reproductive and vegetative proliferative activities.

5.7 Evolution of meristems

Plant form and architecture are tightly linked to the activity of meristems.
Indeterminate meristems have independently evolved in the gametophyte
phase of bryophytes and ferns, and in the sporophyte of vascular plants.
Gametophyte meristems considerably differ from those of sporophyte mer-
istems, because the former are composed of a single stem cell, while the latter
consist of a single stem cell, or a group of cells, overlaying several layers of
proliferative cells (Ambrose & Vasco, 2016; Harrison et al., 2007, 2009;
Langdale, 2008; Sanders, Darrah, & Langdale, 2011). Meristematic activity
is not recognizable in the fossil record of early vascular plants. Furthermore,
fossils at the divergence of vascular plants and bryophytes are missing, which
makes it impossible to decide whether sporophytic indeterminant meristems
have evolved once or multiple times independently. Therefore, all informa-
tion is coming from observations made on extant taxa.
Among the extant representatives of the earliest diverged lineages, mosses,
hornworts and some liverworts have multicellular proliferative sporophytic
regions, which may be—to some extent—homologous to proliferative regions
of the indeterminate meristematic regions of extant vascular plants (Bowman,
2013; Ligrone et al., 2012b; Tomescu et al., 2014). Importantly, hornwort
basal meristems seem to be indeterminate, while moss intercalary meristems
show only transient activity (Bowman, 2013; French & Paolillo, 1975b;
Langdale, 2008; Ligrone et al., 2012a, 2012b; Villarreal & Renzaglia, 2015).
It remains to be seen whether these structures are homologous to one another,
and whether their proliferative activity shares common regulatory activity with
that of the proliferative regions of vascular plant meristems.
The structure of indeterminate meristem activity of sporophyte shoots in
lycophytes and ferns is highly variable, but it usually consists of a single apical
cell, or groups of cells, overlaying a deeper layer of proliferative cells
(Ambrose & Vasco, 2016; Frank et al., 2015; Vasco et al., 2016). Expression
of key genes of the SAM in ferns and lycophytes suggests that their sporo-
phytic meristems are multicellular structures, and their core regulatory
mechanisms may be homologous to that of the flowering plant SAM
(Evkaikina et al., 2017; Friedman, 2011). Nevertheless, transcriptomic
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Title: Kun naiset pelkäävät: Yksinäytöksinen huvinäytelmä

Author: Ville Vuoksinen

Release date: May 10, 2020 [eBook #62088]

Language: Finnish

Credits: Tapio Riikonen

*** START OF THE PROJECT GUTENBERG EBOOK KUN NAISET

PELKÄÄVÄT: YKSINÄYTÖKSINEN HUVINÄYTELMÄ ***
E-text prepared by Tapio Riikonen

KUN NAISET PELKÄÄVÄT

Yksinäytöksinen huvinäytelmä

Kirj.

VILLE AKKANEN [Ville Vuoksinen]

Hämeenlinnassa, Arvi A. Karisto Osakeyhtiö, 1922.

HENKILÖT:

MATTI, isäntä.
KAISU, hänen vaimonsa.
VAPPU, Matin äiti.
HELENA, Matin sisar.
NIITTYNEN, naapuri-mies.
TAHVO KYNTÖLÄ, talonpoika, sulhanen.
MATTILA, puhemies.

Tapaus salokylässä Karjalassa.

NÄYTTÄMÖ: Tavallinen, yksinkertaisesti kalustettu huone. Perällä

ovi; oikealla uuni ja akkuna; vasemmalta vie ovi kamariin.

Matti istuu vasemmalla sijaitsevan pöydän ääressä lukien

sanomalehteä.
Vappu kutoo sukkaa, Kaisu kehrää.

MATTI. Aina niitä varkaitakin vain liikkuu. Joka päivä saa niistä
sanomalehdestä lukea.

KAISU. Missä ne nyt ovat mellastaneet?

 MATTI. Aivan tässä naapuripitäjässä. Erään vaateaitan olivat näet
ihan tyhjentäneet.

VAPPU. Kylläpä ovat ihmisiä!

KAISU. Ei niillä näy säälintunnetta ensinkään olevan; kaiken vaan

vievät.

MATTI. Säälintunnetta! Ketä rosvot säälisivät? Ei niiden

omaatuntoa mikään vaivaa, vaikka tappaisivat ihmisenkin.

VAPPU. Niinhän se taitaa olla. Kunhan vain eivät tänne tulisi.

MATTI. Eipähän niitä vielä täällä ole nähty. — Mutta tuokaahan,

naiset, illallista; kello jo kahdeksatta käy.

VAPPU. Kannahan sinä, Kaisu, ruoka pöytään! Minä saisin tämän

sukan tänä iltana valmiiksi.

KAISU (vähän kiivastuen). Minulla olisi tässä kyllä kehräämistä.

VAPPU. Voithan huomenna kehrätä sen, mitä et tänään ehdi.

KAISU (kiivaasti). Voitte tekin sen sukan valmistaa vasta

huomenna.
Minähän tässä emäntä olen.

MATTI. No… no… Joko taas…

VAPPU (kiivaasti). Sinä emäntä! Niin — sitä sinä olet laulanut aivan
siitä asti, kun Matti-poikani sinut tähän otti. Mutta sen sanon sinulle,
että minä tässä emäntänä heilun niinkauan kun pääni pystyssä
pysyy, enkä alistu kenenkään käskettäväksi.
 MATTI. NO… no… Mitä se äiti taas siinä…

KAISU (melkein itkien). Ja sen minäkin tässä sanon ja sanoisin

vaikka koko kylän kuullen, että niin riitaisaa ja pahansisuista anoppia
ei toista löydy ainakaan tässä pitäjässä ja tuskin toisessakaan.

MATTI (Kaisun puhuessa). No, mitä sinä, Kaisu, taas? Älä nyt
tuossa riitele! Eihän se sillä parane. No… no… Herkeähän siitä!…
Tuokaa se ruoka!…

VAPPU. Vai pahansisuinen! Hahahaa. Kuka se nytkin riidan aloitti?

Minäkö, häh?

HELENA (tulee kamarista). Mikä nyt taas on? Aina te riitelette!

MATTI (Helenalle). Tuo sinä, Helena, illallinen! (Vapulle ja

Kaisulle). Ja te akat, jos ette tukkoa suutanne, niin saatte
uloslähdön. Ja tuokaakin se illallinen nyt juuri pöytään!

KAISU. Kyllä meidän täytyy, Matti, erota talosta. Ei tästä elämästä

tul…

MATTI (keskeyttää kiivaasti). Suu kiinni ja paikalla! — No, ruoka

pöytään! (Vappu ja Kaisu nousevat.)

HELENA. Kyllä minä tuon. (Astuu peräovea kohti. Kopinaa ulkoa.)

MATTI. Kukahan vieras se näin myöhään mahtaa tulla?

NIITTYNEN (astuu sisään). Hyvää iltaa!

MATTI. Iltaa, iltaa! Astukaahan, Niittynen, peremmälle! Mitäs

naapurille kuuluu?
 NIITTYNEN (istuutuu). Onhan niitä kuulumisia. Varkaita ja rosvoja
nyt näillä seuduin majailee.

VAPPU, KAISA ja HELENA (huudahtaen). Täälläkin!

MATTI. Onko niitä nähtykin?

NIITTYNEN. Niin — Ylätalon Maija se päivällä marjassa

käydessään ne tuolla salon puolella oli nähnyt. Pitkiä miehen
roikaleita kuuluivat olleen, ja heti heidät huomattuaan Maija pyyhälti
kotiin minkä ennätti.

MATTI. Vai niin. Vai jo tännekin alkavat ilmestyä. Tässä lehdessä

kerrotaan myöskin naapuripitäjällä rosvojen mellastaneen. Eiköhän
vain liene samoja roistoja?

NIITTYNEN. Samaa mekin kotona arvelimme. Niitä myöskin oli

ollut kaksi, niinkuin Maijakin sanoi nähneensä. (Hetkisen perästä). Ja
kun se meidän torppa siellä on syrjässä toisista taloista, eikä meillä
ole miehiä muita kuin minä, niin tulin tänne pyytämään isäntää
vaikkapa täksi yöksi kanssani sinne meille. Eivät sitten ainakaan
uskalla sisään tunkeutua, ja jos tulisivatkin, niin selkään antaisimme.

MATTI. Hm, pitäisihän sitä naapuria senverran auttaa.

VAPPU. Mutta jos ne sillä aikaa tulevatkin rosvoilemaan meille.

Mitä me heille mahdamme!

HELENA. Niin, parasta taitaa olla, että jäät, Matti, kotiin.

KAISU. Jää vaan kotiin, jää.

 NIITTYNEN. Eipä ne taida näin kylän laidassa näyttäytyä. Onhan
tässä lähellä muitakin taloja. Mutta toista on meillä; ainakin puoli
kilometriä on lähimpään naapuriin, teille nimittäin, ja vielä on
metsääkin välillä. Sinne vorot hyvästi uskaltavat tunkeutua. Teille ne
eivät mitään pahaa tule tekemään.

MATTI. Niinhän se on. Eivät ne tänne uskalla. Pitäähän aina

naapuria auttaa. Mutta syödään ensin illallinen.

NIITTYNEN. Syödään illallinen siellä meillä. Ne naiset taitavat

kotona olla peloissaan, jos viivymme.

MATTI. NO lähdetään sitten. (Nousevat.)

VAPPU. Voi voi sentään! Jos ne vaan hyökkäävät sillä aikaa meidän
päällemme.

MATTI. Älkää pelätkö! Eivät ne tänne uskalla tulla.

HELENA. Niin. Eihän nuo nyt toki näin kylän laitaan. (Matti ja
Niittynen lähtevät.)

KAISU. Ja kyllähän minussakin jonkun verran miehelle vastusta

riittää, vaikka olenkin nainen.

VAPPU. Tuskinpa vain.

KAISU. Kyllä! Ja jos ne, senkin maailman tasaajat, vain tänne

meille vieraiksi tulevat, niin selkään mekin niille annamme. Kyllä
minä toisen miehen osalleni otan.

HELENA. Kyllä me taas äidin kanssa toisen kuritamme.

 VAPPU. Ei ole minusta vanhasta enää mihinkään. Mutta autanhan
minä teitä, jos ei muussa niin huutamisessa.

KAISU. Me emme huudakaan. Eikä teissä taitaisi olla auttajaa

huutamiseenkaan.

HELENA. Huh, kyllä olisi sentään kamalaa, jos meidän täytyisi

rosvojen kanssa tappelemaan käydä. (Hetken perästä.) Pitäisihän se
illallinenkin syödä. (Astuu peräovea kohti.)

KAISU. Ei minulla ainakaan nälkä ole. Niitä varkaita tässä

ajattelen.

VAPPU. Ei ole minullakaan. Syö sinä, Helena, jos haluat!

HELENA. En minäkään sitten kehtaa syödä. Syödään

myöhemmällä.

KAISU. Kyllä lehmille ainakin illallinen kelpaa. Ja ne vaateaitatkin

pitää lukita. (Poistuu.)

HELENA (katselee sanomalehteä). Mistä asiasta te äsken riitelitte?

VAPPU. Eihän siihen paljoa tarvita, kun Kaisu riidan aloittaa. Siitä
emännyydestä se vähän kiukustui. Mutta niinkuin ennenkin, hän
nytkin pian leppyi.

HELENA. Etkö vaan lie, äiti, taas häntä härnännyt? Eihän olekaan
ihme, jos hän sattuu suuttumaan.

VAPPU. Minä härnännyt! Hänhän härnää ja komentelee minua,

ikäänkuin olisikin aivan emäntänä talossa.
 HELENA. Saisithan, äiti, hänelle emännyyden luovuttaa, kun hän
niin tahtoo. Melkein joka päivä teillä on siitä riitaa.

VAPPU. Niin — olen sitä kyllä ajatellut. Kuitenkin aion emännyyden

pitää ainakin siihen asti, kun sinä miehelään joudut. En henno antaa
sinua hänen käskettäväkseen.

HELENA (arasti). Ei taida äiti vielä tietääkään, että minulla jo

melkein onkin sulhanen.

VAPPU. Sinulla jo sulhanen! Kuka se on?

HELENA. Eräs Vaaralan kylän poika.

VAPPU. Vai niin. Jo minä sitä vähän olen sivustapäin kuullut

kerrottavan. Hm, ei sinulla vielä olisi miehelään kiirettä, vasta
keväällä rippikoulusi päätit. Onko se edes talonpoikakaan?

HELENA. Rikkaan talon poika onkin. Hän on vielä vanhempiensa

ainoa poika.

VAPPU. Vai niin. Mikä hänen nimensä on?

HELENA. Tahvo Kyntölä.

VAPPU (hämmästyen). Siitäkö talosta se on, jossa on suuri

kivinavettakin?

HELENA. Juuri sen talon poika.

VAPPU. Hm, rikas on sulhasesi. Tuskin vain huolinee näin köyhän

talon tyttärestä. — Pettää sinut.
 HELENA. Saattepa nähdä. Tällä viikolla jonakin päivänä hän tulee
tuomaan kihlat, ja ensi viikolla menemme jo pappilaan kuulutuksille.

VAPPU. Sinusta tuleekin hyvä miniä apellesi ja anopillesi. Et

olekaan sellainen riitapukari kuin tuo Kaisu. — Kyllähän Kaisukin
varma ja ahkera on, mutta vähän kivakka se on luonteeltaan.

HELENA. Tahvon äiti onkin jo kuollut; anopista siis ei ole mitään

vastusta.

VAPPU. Vai jo kuollut. Sitten sinä pääsetkin emännäksi heti.

KAISU (rientää hengästyneenä sisään). Voi voi tätä kauhistusta!

Nyt ne ryövärit istuvat tuolla ruispellon pientareella.

VAPPU ja HELENA (säikähtäen). Mitä! Ruispellolla?

KAISU. Niin. Kun tulin navetasta ja satuin katsahtamaan tuonne

polulle päin (viittaa ikkunaan), huomasin kaksi miestä, jotka istuivat
pientareella ja katselivat tännepäin. (Kaikki puhuvat hätääntyneinä ja
kurkistelevat akkunasta.)

VAPPU. Voi voi! Ne varmaankin tuumivat, mikä olisi viisaampaa:

tunkeutua ensin vaateaittoihin, vai rynnätä suoraan tupaan.

KAISU. Nyt ne vievät sieltä aitasta minun puolisilkkisen huivini,

jonka
Matti vasta keväällä osti!

VAPPU. Ja minun uuden pässinnahka-turkkini, jota vain yhden

talven olen pitänyt!

HELENA. Ja minun pitsireunuksisen alushameeni!

 VAPPU (kiivaasti). Mutta miksi sinä, Kaisu, et niitä huoneita
lukinnut?

KAISU. Lukitsinhan minä ne. — Mutta eihän se mitään auta.

VAPPU. Ei auta! Minkätähden?

KAISU. Sentähden, että varkailla on näet sellaiset avaimet, joilla

pääsee vaikka kirkkoonkin.

HELENA. Mistä sinä tiedät, että näillä sellaiset on?

KAISU. Onhan ne kaikilla pitkäkyntisillä.

VAPPU. Voi voi, mitä me nyt teemme?

KAISU. Niin — mikä nyt neuvoksi tulee? Ja kun ne näkevät, että

me olemme kaikki naisia, niin uskaltavat aivan hyvin rynnätä
vaateaittoihin ja tupaankin, — ja jos vielä tappavatkin meidät. Eihän
niiden omaatuntoa mikään vaivaa.

HELENA. Voi voi! Kun meistä nyt vaikka yksikin olisi mies!

KAISU. Niin — kun olisi! Mutta mistä sen tähän miehen tempaa.
Kuka sitä uskaltaa naapuristakaan mennä hakemaan.

VAPPU. Eiköhän sentään kova huuto auttaisi!

KAISU. Huutaminenko? Silloinhan me juuri olisimme kuoleman

omat. Jos rosvot vaan kuulisivat, että me apua huudamme, niin
paikalla ne sisään hyökkäisivät ja tappaisivat meidät.

VAPPU. Voi tätä kauhistuksen iltaa!

 KAISU. Mutta nyt pälkähti oiva tuuma päähäni. Minä rupeankin
mieheksi.

VAPPU ja HELENA. Mitä?

KAISU. Minä rupean mieheksi. Puen Matin vaatteet, jotka ovat

tuolla kamarissa, päälleni, niin — silloinhan olenkin jo mies. Ainakin
siltä näytän, ja eihän muuta tarvitakaan. Eivät rosvot sitten uskalla
pihallekaan tulla, kun näkevät tuvassa olevan yhden
mieshenkilönkin.

HELENA. Mainio tuuma. Pukeudu nyt sukkelaan; kohta voivat olla

jo täällä.

KAISU. Niin teenkin. (Poistuu kamariin. Vappu ja Helena

kurkistelevat ikkunasta ja kuiskailevat keskenään.)

KAISU (kamarista). Haha. Hyvä tuuma se on. Eivät varkaat

uskallakaan tulla kaivamaan ja varastamaan. — Mutta kyllä nämä
Matin housut ovat jotensakin avarat. Leveäthän ne tosin pitää
kyntömiehen lantiot ollakin. — Kuules, Helena, kierrä sitä tulta vähän
suuremmaksi! Nähkööt pitkäkyntisetkin, ettemme täällä pimeässä
piileile!

HELENA. Niin oikein. (Kiertää tulta lampussa suuremmaksi.)

KAISU (kamarista). Onhan tässä sarkaviitassakin tilaa. Jykevät

ovat hartiat Matilla; suorat ja syvät ovat sentään ojatkin Takakorven
suolla. Ja voi sitä Matin päätä, kun se on suuri! Sanotaan: suuri pää,
vähän mieltä, mutta mitä turpeenpuskijalla tarvitsee enempää
viisautta ollakaan, kun osaa vain pitää peltonsa kunnossa ja kykenee
raivaamaan toista korpeen ja suohon. — Eihän niitä rosvoja vielä
näy?

HELENA. Tuolla nuo veräjän luona seisovat.

KAISU (kamarista). Nyt olenkin valmis. (Tulee kamarista,

kannatellen toisella kädellään leveitä housuja, pitkä sarkatakki yllään
ja korviin asti painunut hattureuhka päässä.) No, näytänkö minä
mieheltä? (Vappu ja Helena nauravat.)

VAPPU. Hahhahaa, minkänäköinen sinä olet!

HELENA. Mieheltä näytät muuten, mutta ei vain ole partaa nenän

alla.

KAISU. Niin — eihän se Luoja ole naisille partaa suonut, mutta

pianhan sen saapi. (Ottaa liedestä hiilen, jolla vetää viikset itselleen.)

VAPPU. Kyllä nyt jo aivan mieheltä näytät. — Kun olisi sinulla

miehen luontokin, niin ei olisi mitään hätää.

HELENA (katsoen ikkunaan). Voi voi, nyt ne avaavat veräjän ja

lähtevät tännepäin.

VAPPU. Kyllä me nyt olemme hukassa.

KAISU. Hätä keinon keksii: pannaanpa tämä mukurapää-puntari

pöydälle. Ja ihme on, jos ne eivät miestä ja puntaria nähdessään
säikähdä ja pakoon pötki. (Tuo ovensuusta puntarin ja koettaa
asettaa sitä pöydälle pystyyn, saamatta kuitenkaan.) Ole sitten
pitkälläsi, kun et pystyssä pysy!
 HELENA. Nyt ne ovat jo tuossa saunan luona. Voi kun se Mattikin
meni sinne Niittyselle.

KAISU. Niin, eihän ne miehet sitä usko, mitä me naiset sanomme,

vaikkemme ole heitä yhtään huonompia.

VAPPU. Voi voi voi!

HELENA. Kyllä tässä sentään yksi oikea mies olisi hyvään

tarpeeseen.

KAISU. Niin — olisihan se! Eihän niiden rosvojen kanssa naiset

oikein uskalla tappelemaan käydä.

VAPPU (ilkkuen). No, mitä sinä pelkäät? Mieshän sinä nyt oletkin.
Ja kyllähän Kaisu yhden miehen kurittaa, koskapa tässä viime viikolla
hyvästi riitti Ylätalon Maijalle ja Kustaalan Mantalle molemmille
tukkanuottasilla ollessa. Tokihan kaksi naista nyt yhtä miestä vastaa,
ja…

KAISU (keskeyttäen). Mitä! Minäkö olen milloin tapellut?

Todistakaa se! Olenko minä sellainen? Häh! Olenko minä
tappelunhaluinen? (Lähenee uhkaavasti Vappua.) Itsehän tuossa
aina riitaa haastatte!

VAPPU (Kaisun puhuessa). Niin, niin. Näinhän sen aivan selvästi

saunan ikkunasta. Mitä? Minäkö riitaa haas…

HELENA (keskeyttäen). Heretkää siitä jo! Nyt ne ovat tuossa

pihalla. Voi voi! Tänne tupaan näkyvät tulevan. (Naiset siirtyvät
pelokkaina pöydän luo.)
 VAPPU (osoittaen ovea). Mutta voi voi voi! Ovensäppihän on auki!
Mikset pannut sitä kiinni, Kaisu?

KAISU. Nyt sekin on taas minun syyni. Olisitte itse pannut!

VAPPU. Sinähän viimeksi ovensuussa kävit puntaria hakemassa.

KAISU. Minä en ainakaan uskalla mennä sitä kiinni panemaan.

VAPPU ja HELENA. Voi voi voi!

KAISU. Voi voi! Nyt ne tappavat meidät. Voi mihin minä nyt
joudun! Minun päällenihän ne ensiksi hyökkäävät, minä kun olen
miehen vaatteissa. Voi voi, miksi rupesinkaan mieheksi.

HELENA (tarttuen pöydällä olevaan juoma-astiaan). Tästä minä

heille annan! (Kopinaa ulkoa. Tahvo ja Mattila astuvat sisään.
Samassa syntyy kuvaamaton meteli. Helena heittää juoma-astiassa
olevan piimän Tahvon päälle. Vappu ja Kaisu hyökkäävät kädet ojona
tulijain kimppuun. Helena huitoo juoma-astialla, Vappu käsillään,
Kaisu raapii Mattilaa, heittää hänen lakkinsa maahan ja tarttuu
lopuksi hänen tukkaansa. Tulijat hämmästyksissään koettavat
käsillään työntää naisia etemmäksi. Kaikki tapahtuu nopeasti.)

KAISU (ollen Mattilan tukassa). Vai, senkin roistot, hyökkäätte

turvattomien naisten ja leskien kimppuun. Kyllä minä teille näytän!
Tuntuuko päänahassasi? Häh? Mitä sanot? Tuntuuko?

MATTILA (huutaa rimpuillen). Mitä peliä tämä on? Älä hyvä mies
revi minun tukkaani! Hullujako te olette?

TAHVO (käsillään suojellen päätänsä). Mitä tämä merkitsee?

Tällälaillako meitä vastaanotetaan! Mitä sinä huidot, Helena?
 HELENA (hämmästyen). Tahvo!

VAPPU (hämmästyen). Sulhaset!

KAISU (jättää Mattilan rauhaan). Mitä te sanoitte?

HELENA (hämillään). Niin… Kaisu, me vähän erehdyimme. Nämä

ovatkin kosijoita.

KAISU (ihmetellen). Vai kosijoita!

MATTILA. Kaisu! (Kaisulle.) Ettekö te olekaan mies?

KAISU (häpeissään). E-enhän minä.

MATTILA. Isännäksi minä teitä luulin. Mutta miksi te olette miehen

pukimissa?

KAISU (häpeissään). Niin — ne varkaat —

TAHVO. Luulitteko te meitä varkaiksi?

HELENA. Niin, tuolla metsässä olivat päivällä nähneet pari

epäilyttävän näköistä miestä, ja tämän takia meni Matti-veljeni
Niittylään yöksi; nämä näet pelkäsivät rosvojen sinne tunkeutuvan.
Mutta kun Kaisu navetasta tultuaan huomasi teidät tuolla ruispellon
pientareella istumassa, luulimme teitä samoiksi roistoiksi. Ja niitä
muka peloittaakseen pukeutui Kaisu Matin vaatteisiin.

MATTILA. Jo minä sitä epäilin, mitenkä mies voi toisen miehen

tukkaan tarttua; sehän on naisten tapaista.

KAISU. Naurakaa vaan! Mutta kyllä te taisitte hyvän tukkapöllyn

saada?
 MATTILA. Vieläkin tuntuu se päänahassani. (Nauravat.)

KAISU. Kyllähän minä teitä jotenkin kovakouraisesti pitelin,

mutta… (katsahtaa ikkunaan) voi voi, nythän ne vasta oikeat rosvot
tulevatkin. Katsokaa, kuinka seipäät olalla juoksevat! (Katsovat
ikkunaan. Naiset voivoitavat.)

VAPPU ja HELENA. Voi voi! Nyt ne tulevat! Puolustakaa meitä!

TAHVO. Kyllä me heidät kuritamme.

MATTILA. Menkää te vain kamariin, älkääkä huutako!

HELENA. Mutta jos ne tappavat sinut, Tahvo?

TAHVO (työntäen Helenaa hellästi kamariin). Älä pelkää! Ei ne

meille mitään mahda. (Naiset menevät kamariin. Tahvo ja Mattila
asettuvat molemmin puolin ovea.)

TAHVO. Minä otan vastaan ensiksitulevan, sinä saat pitää huolen

jälkimäisestä.

MATTILA. Tuossa ne jo ovatkin. (Matti ja Niittynen ryntäävät

samassa sisään. Tahvo tarttuu Matin rintapieliin, Mattila Niittysen.
Riuhtovat toisiaan ja puhuvat yhtaikaa. Naiset voivottavat metelin
aikana kamarissa.)

MATTI. Senkin roistot, vielä teette vastarintaa! Ulos ei ole

pyrkimistäkään. Niittynen, pidä puolesi!

NIITTYNEN. Äh! Annan heille oikein miehen kädestä.

TAHVO. Moiset rosvot! Nyt ette vähällä pääsekään. Naiset, tuokaa

nuoria! Huomenna viedään teidät nimismiehen luo ja ylihuomenna
kuritushuoneeseen!

MATTILA. Kiikkiin jouduitte, pitkäkyntiset!

MATTI. Mitä! Minut kuritushuoneeseen! Kyllä minä teille näytän!

HELENA (kurkistaa kamarin ovelta). Tässä on köyttä.

(Hämmästyen.) Mitä!
Isäntähän se onkin ja Niittynen! (Ottaa Tahvoa käsipuolesta.)
Hellittäkää jo! Hän on isäntä. (Miehet eroavat.)

TAHVO (hämmästyen). Isäntä!

MATTI. Mutta mitä miehiä te oikeastaan olette, kun noin vain

uskallatte rynnätä isännän päälle ja vielä hänen omassa tuvassaan?

TAHVO. Luu… luulimme teitä varkaiksi.

MATTI. Varkaiksi?

HELENA. Tulehan, Matti, tänne, niin selitän asian. (Poistuvat uunin

luo. Kuiskailevat keskenään.)

TAHVO. Nyt me vasta huonosti teimme, Mattila. Mitähän isäntä

sanoo?

MATTILA. Huonosti teimme. Mutta emmehän tienneet, että hän oli

isäntä.

TAHVO. Tokkopa asiamme nyt rupeavat luonnistamaan kun

tällaisen kepposen teimme.

MATTILA. Kyllä. Siitä saat olla varma.

 MATTI (kääntyen vieraiden puoleen). Vai niin. Te luulitte meitä
varkaiksi niinkuin mekin teitä. Sattuuhan sitä erehtymään. — Vai
kosijoita olettekin. Käykäähän peremmälle! (Istuutuvat.) Pannaanpa
tupakaksi. (Ottaa pöytälaatikosta savukelaatikon ja tarjoaa jokaiselle.
Tupakoivat.)

KAISU (huutaen kamarista). Joko ne rosvot ovat köysissä?

HELENA (nauraen). Jo ovat. Tulkaa katsomaan! (Kaisu ja Vappu

astuvat arkoina tupaan.)

KAISU. Aha! Joko jou… (Hämmästyy.) Ka, Mattihan se onkin!

MATTI (ihmeissään). Mitä "komediaa" sinä, Kaisu, olet pelannut?

KAISU (häpeissään). Niin — ne varkaat… (Pujahtaa kamariin.

Vappu jää tupaan.)

MATTI. Mikä hätä teillä oli varkaiden tähden? Olihan teidän

turvananne tämä kosiomies puhemiehineen. (Katsahtaa Tahvoon.)
Mutta miten te olette noin piimässä?

TAHVO. He luulivat meitäkin varkaiksi. Kun tulimme tupaan ja

avasimme oven, ryntäsivät naiset meidän päällemme. Minä sain
piimätuopin sisällyksen niskaani (taputtaen Mattilaa olalle), mutta
Mattila se vasta pulassa oli: Kaisu näet riippui molemmin käsin
hänen tukassaan, niin ettei mies-poloinen joutanut oikein
huutamaankaan. (Nauravat.)

MATTI. Niin — ne naiset ovat naisia.

HELENA (ottaen pyyheliinan). Saanko puhdistaa, Tahvo, takkisi?

 TAHVO. Miksei. — Ethän minua enää koskaan piimää. —
(Kuiskailevat keskenään. Helena puhdistaa takin.)

NIITTYNEN (Mattilalle), Me taas kun näimme teidän tulevan tuolta

metsän läpi ja suuntaavan kulkunne tännepäin, aloimme seipäiden
kanssa hiipiä jälestänne. Ja kun huomasimme, että astuitte tupaan,
lähdimme heti juoksujalkaa perästä ja…

MATTI (keskeyttäen, Mattilalle). Hahahaa! Te taas luulitte meitä

rosvoiksi ja otitte oikein miehen kourilla vastaan.

MATTILA. Aivan niin. Ja se olikin pulskempi yhteenotto kun akkain

kanssa.

MATTI. Minäkin kouristin jo aivan hartiavoimallani. (Kaisu astuu

kamarista häpeissään, entisessä naisenpuvussaan tupaan.)

MATTI (nauraen). Hahahaa. Tuo Kaisuhan minua vähän naurattaa.

Kun hätä tulee, niin pukeutuu miehen vaatteisiin ja kuitenkin sitten
tukassa riippuu. Etkö sinä, Kaisu, jo raamatustakin tiedä, että eihän
nainen voi milloinkaan olla mies, vaan ainoastaan miehen apu.
(Miehet nauravat.)

KAISU (puolustellen). Täytyihän sitä hädässä keksiä jos vaikka

mitä.

MATTI. Niin — jokaisellahan meillä on ollut huolta ja hommaa

varkaiden takia, mutta kuitenkaan ei niistä ole tietoakaan.

NIITTYNEN. Siihen Maijaankaan ei ole mitään luottamista. Jos ne

hänen salolla näkemänsä miehet olivatkin vaikka metsästäjiä.

MATTILA. Meitä vastaan tulikin kaksi miestä, joilla oli pyssyt olalla.
 MATTI. No niin! Nehän se Maija oli nähnytkin.

KAISU. Harvoinhan ne Maijan sanat uskottavia ovat.

MATTI (Mattilalle). Mutta miksi te tulitte jalkaisin? Missä teidän

hevosenne on?

MATTILA. Sen jätimme Mäkelän lautamiehelle ja tulimme tästä

metsän läpi suoraan.

TAHVO (kuiskaa Mattilalle). Esitä nyt se asia!

MATTILA. Niin — tuota. Pitäisihän se asiakin toimittaa. — Kosijoita

olemme, niinkuin tiedätte. Tämä Kyntölän Tahvohan se on
kainalokanaistansa etsimässä, ja tästä talosta on hän sen löytänyt.
Iso on Tahvolla koti: kaksitoista pulskaa lehmää seisoo suuressa
kivinavetassa, kolme on hevosta; paljon on kylvetty ruistakin:
kokonaista kah…

MATTI (keskeyttäen). No, no, no! Kyllä minä tiedän, mitä

Kyntölässä kylvetään ja mitä niitetään. Iso on talo. (Tahvolle.)
Mieluinen olet vävy meille, Tahvo. (Helenalle.) Kelpaako Helenalle
kihlat?

HELENA. Kyllä kelpaa.

TAHVO. No, katsellaanpa sitten näitä kihlasia. (Vetää

povitaskustaan paperikääröjä.)

MATTI. Ja nyt, Kaisu ja äiti, tuokaapa viimeinkin se illallinen!

Väliverho.
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