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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.
https://doi.org/10.1016/bs.ctdb.2018.11.005
2 P
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enyi et al.

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
eter Sz€
ov
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
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€
ov
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;
Discovering Diverse Content Through
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well matured. He could see no reason why the business should be
precipitated upon the House—a proper delay would not show any
want of respect to the President, as some gentlemen think. Would it
be more respectful that an answer should be sent by this House,
which, for want of time, had not been sufficiently considered?
Certainly not. Far more so will it appear that after mature
deliberation the members are unanimous in their answer. I therefore
think the object of respect which the gentleman from North Carolina
has in view will be completely answered by the delay.
Gentlemen talk about precedent. I am ashamed to hear them. There
may be no precedent on the subject. But are we always to act by
precedent? There is scarcely a circumstance occurs in this House but
what is different from any that was before it. The President's
Addresses to this House are always different. They relate to the
circumstances of things that are, have been, and may be. Then, to
talk of precedents where things cannot be alike, is to trammel men
down by rules which would be injurious in the issue.
The Message of the President respecting the French Colors had been
referred to. If gentlemen were then wrong, is that a reason why
they should continue to act wrong? But this circumstance materially
differs from that. That was merely an expression of sentiment, which
could at once be determined, but this of sentiment, accompanied
with deep and solemn reflection—it is so interwoven with the politics
of the country as to require great circumspection. I hope gentlemen
will not go into it until they are properly prepared. I wish to pay all
possible respect to the Chief Magistrate, and cannot prove it better
than by a sincere desire for an unanimous vote to the answer, which
is only to be obtained by proper deliberation; and thus let him
depart from his office with credit, and the enjoyment of our best
wishes in his retirement.
The question for postponing the unfinished business to take up this
report was then put and negatived—43 to 31.

Wednesday, December 14.

Thomas Henderson, from New Jersey, and Thomas Hartley, from
Pennsylvania, appeared and took their seats in the House.

Reporting of the Debates.

Mr. W. Smith moved for the order of the day on the petitions of
Thomas Lloyd and Thomas Carpenter, whereupon the House
resolved itself into a Committee of the Whole, when, having read the
report of the committee to whom it was referred,
Mr. Macon wished some gentleman who was in that committee,
would be so good as to inform the House what would be the
probable expense, and for what reason the House should go into the
business. He thought the expense altogether unnecessary, whatever
it may be.
If the debates of this House were to be printed, and four or five
copies given to each member, they would employ all the mails of the
United States. He also adverted to the attempt at the last session to
introduce a stenographer into the House, which failed.
Mr. Smith informed the gentleman that Mr. Lloyd's estimate of the
expenses is, that he will supply the House with his reports at the
rate of three cents per half sheet. His calculation is that he can
supply the members at the expense of about $1,600 for the session.
With respect to the gentleman's reference to last session, this was
materially different from that: that motion was to make the person
an officer of this House, and at an expense much greater. He
thought this attempt would be of great use to the House. Regular
and accurate information of the debates in the House would be a
very desirable thing; he therefore hoped the resolution would prove
agreeable to the House.
Mr. Williams said, that the House need not go into unnecessary
expense: the members were now furnished, morning and evening,
with newspapers, which contained the debates; then why should the
House wish for more? If one person in particular has the sale of his
debates to this House, will it not destroy the advantages any other
can derive from it? We ought not to encourage an undertaking of
this kind, but let us encourage any gentleman to come here and take
down the debates. Last year they were taken down very accurately
and dispersed throughout the Union.
By passing this resolution you will destroy the use of the privilege to
any other than the person favored by this House. Why give one a
privilege more than another? He observed, it had been common to
give gentlemen the privilege to come into the House and take down
the debates, which had been, last year, delivered time enough to
give satisfaction to the members.
Mr. Thatcher said, he should wish for information from the committee
how many persons there were to publish debates, as he understood
there were several, and the members were to supply themselves
from whom they pleased. He should likewise wish for information,
how many each member was to have to amount to the value of
$1,600.
Mr. W. Smith said, there had been petitions received from only two
persons—Thomas Lloyd and Thomas Carpenter. They intended, each
of them, to publish the debates. There might be others; he knew
not. There was no intention of giving any one a preference—
gentlemen could subscribe for that they approved of most. At the
calculation of Mr. Lloyd the members would have five copies each for
the $1,600.
Mr. W. Lyman said, the question was, whether the House would incur
the expense of $1,600 to supply the members with copies or not?
He thought there was no need of the expense. If the House do not
think proper to furnish the members, they can supply themselves. A
publication of them is going on at present, and many gentlemen had
subscribed to it already.
Mr. Dearborn did not think that $1,600 thus laid out would be
expended to the best possible advantage. From the number of
persons which we see here daily taking down debates, he thought
we might expect to see a good report of the occurrences in the
House. There was a book going about for subscriptions, which
appeared to be well encouraged; he saw many of the members'
names in it. He thought that, by a plan like that, the reports may be
as accurately taken as we may have any reason to expect if the
House incurs this expense.
Mr. Nicholas observed, that members were now served with three
newspapers. He thought to vote for this resolution on account of
obtaining a more full and complete report than was to be had in the
newspapers; thus it would supersede the necessity of taking so
many papers. He thought this plan more useful to the members, and
generally of more advantage to their constituents, as they could
disperse those debates where otherwise they would not be seen.
Mr. Thatcher said, if the object of the motion was to supersede the
receiving of newspapers, he certainly should vote against it. He did
not consider the main reason why members were served with the
newspapers was, that they may obtain the debates. No. He thought
it more important, in their stations, that they should know the
occurrences of the day from the various parts of the United States as
well as from foreign nations. Though he might favor an undertaking
of this kind, yet he would give preference to a newspaper, if they
were to have the one without the other.
Mr. Heath did not wish that the members, being furnished with
debates agreeably to the motion, should supersede the receiving of
newspapers, yet he should vote for it. Gentlemen had said the
debates were taken more correctly last session than before, yet he
had heard a whisper which was going from North to South, that our
debates are not represented impartially. He wished the House and
the people to be furnished with a true report; such a thing would be
very useful: however, he did not wish to encourage a monopoly to
those two persons. No. He would wish to give an equal chance to all
who choose to come and take them. Shall we repress truth? I hope
not; but disseminate it as much as possible. Last session, when I
was, under the act of God's providence, prevented from attending
the House, a member sent for a gentleman from Virginia, who was
to act as stenographer, with whom the House and a printer in this
city were to combine. Warm debates ensued on the propriety of the
measure, and the gentleman returned home after the motion was
negatived. I hope gentlemen will not grudge 1,600 dollars towards
the support of truth. What we see now in the newspapers is taken
from the memory, and not by a stenographer. The people will thank
you that you have taken means to investigate truth. If any
gentleman can point out a better mode to obtain this object, I hope
he will do it that it may be adopted; till then I shall support the
resolution.
Mr. Sherburne did not think, with the gentleman last up, that the
interest of the country was concerned; the only thing they were
concerned in was the payment of the money. The printing of this
work did not depend on the motion of this House. Whether the
House adopt it or not, the book will be published. It is a matter of
private interest; a speculation in the adventurer, like other
publications. The question, he conceived, meant only this: Should
the members be supplied with these pamphlets at the expense of
the public, or should they put their hands in their own pockets and
pay for them individually? He thought the House had no greater
reasons to supply the members with this work than other
publications; they might as well be furnished with the works of Peter
Porcupine, or the Rights of Man, at the public expense.
Mr. W. Smith said, the gentleman was mistaken with respect to the
work going on, whether supported by the House or not. It was true
as it respected the work proposed by Mr. Carpenter; but, with
respect to Mr. Lloyd, he declared he could not undertake it, except
the House would subscribe for five copies for each member.
Mr. Swanwick considered the question to be to this effect: whether
the debates be under the sanction of the House or not? A gentleman
had said, it will be a great service to the public to have a correct
statement of the debates. I think the most likely way to obtain it
correctly is to let it rest on the footing of private industry. We have a
work, entitled The Senator, in circulation. I have no doubt but the
publisher will find good account in the undertaking. Why should the
House trouble itself to sanction any particular work? Gentlemen
would then have enough to do every morning in putting the debates
to rights before they were published, as they would be pledged to
the accuracy of the reports. I never heard that, in the British House
of Commons or Lords, such a motion was ever made, nor have I
ever heard of such in any other country; then why should we give
our sanction and incur a responsibility for the accuracy of it. He said
he should vote against the motion, but would encourage such a
work while it rested on the footing of private adventure.
Mr. Thatcher said, he differed much from the gentleman last up, as it
respected the responsibility of the House on such a publication. He
thought it might as well be said, that because there had been a
resolution for the Clerk to furnish the members of this House with
three newspapers, the House was responsible for the truth of what
those newspapers contained; if it was so, he should erase his name
from his supply of them, as he thought, in general, they contained
more lies than truth. Two considerations might recommend the
resolution. It would encourage the undertaking, and also add to the
stock of public information: on either of these, he would give it his
assent. Soon after he came into the city, a paper was handed him
with proposals for a publication of this kind (The Senator). He, with
pleasure, subscribed to its support; as to general information, that
was given already by newspapers, and though each member was to
be supplied with five copies, yet very few would fall into hands
where the newspapers did not reach. The work would go forward at
any rate. If he thought the work depended on the motion, he should
rejoice to give his vote toward its aid. On the question being put,
only nineteen gentlemen voted in favor of the resolution; it was
therefore negatived.
The committee then rose, and the House took up the resolution.
Mr. Thatcher observed, the question was put while he was
inattentive: he wished it to lie over till to-morrow.
Mr. Giles wished to indulge the gentleman in his desire.
Mr. Thatcher then moved for the vote of the House, whether the
report of the Committee of the Whole be postponed. Twenty-four
members only appearing for the postponement, it was negatived.
The question was then put, whether the House agreed to the report
of the Committee of the Whole and disagreed with the report of the
select committee; which appeared in the affirmative. The motion
was therefore lost.

Address to the President.

The House again resolved itself into a Committee of the Whole on

the Answer to the President's Address; when the Answer reported by
the select committee was read by the Clerk, and then in paragraphs
by the Chairman, which is as follows:
Sir: The House of Representatives have attended to your
communication respecting the state of our country, with all the
sensibility that the contemplation of the subject and a sense of duty
can inspire.
We are gratified by the information that measures calculated to
ensure a continuance of the friendship of the Indians, and to
maintain the tranquillity of the interior frontier, have been adopted;
and we indulge the hope that these, by impressing the Indian tribes
with more correct conceptions of the justice, as well as power of the
United States, will be attended with success.
While we notice, with satisfaction, the steps that you have taken in
pursuance of the late treaties with several foreign nations, the
liberation of our citizens who were prisoners at Algiers is a subject of
peculiar felicitation. We shall cheerfully co-operate in any further
measures that shall appear, on consideration, to be requisite.
We have ever concurred with you in the most sincere and uniform
disposition to preserve our neutral relations inviolate; and it is, of
course, with anxiety and deep regret we hear that any interruption
of our harmony with the French Republic has occurred; for we feel
with you and with our constituents the cordial and unabated wish to
maintain a perfectly friendly understanding with that nation. Your
endeavors to fulfil that wish, (and by all honorable means to
preserve peace, and to restore that harmony and affection which
have heretofore so happily subsisted between the French Republic
and the United States,) cannot fail, therefore, to interest our
attention. And while we participate in the full reliance you have
expressed on the patriotism, self-respect, and fortitude of our
countrymen, we cherish the pleasing hope that a mutual spirit of
justice and moderation on the part of the Republic will ensure the
success of your perseverance.
The various subjects of your communication will, respectively, meet
with the attention that is due to their importance.
When we advert to the internal situation of the United States, we
deem it equally natural and becoming to compare the tranquil
prosperity of the citizens with the period immediately antecedent to
the operation of the Government, and to contrast it with the
calamities in which the state of war still involves several of the
European nations, as the reflections deduced from both tend to
justify, as well as to excite, a warmer admiration of our free
constitution, and to exalt our minds to a more fervent and grateful
sense of piety towards Almighty God for the beneficence of His
providence, by which its administration has been hitherto so
remarkably distinguished.
And while we entertain a grateful conviction that your wise, firm,
and patriotic Administration has been signally conducive to the
success of the present form of Government, we cannot forbear to
express the deep sensations of regret with which we contemplate
your intended retirement from office.
As no other suitable occasion may occur, we cannot suffer the
present to pass without attempting to disclose some of the emotions
which it cannot fail to awaken.
The gratitude and admiration of your countrymen are still drawn to
the recollection of those resplendent virtues and talents which were
so eminently instrumental to the achievement of the Revolution, and
of which that glorious event will ever be the memorial. Your
obedience to the voice of duty and your country, when you quitted
reluctantly a second time the retreat you had chosen, and first
accepted the Presidency, afforded a new proof of the devotedness of
your zeal in its service, and an earnest of the patriotism and success
which have characterized your Administration. As the grateful
confidence of the citizens in the virtues of their Chief Magistrate has
essentially contributed to that success, we persuade ourselves that
the millions whom we represent participate with us in the anxious
solicitude of the present occasion.
Yet we cannot be unmindful that your moderation and magnanimity,
twice displayed by retiring from your exalted stations, afford
examples no less rare and instructive to mankind than valuable to a
Republic.
Although we are sensible that this event, of itself, completes the
lustre of a character already conspicuously unrivalled by the
coincidence of virtue, talents, success, and public estimation, yet we
conceive that we owe it to you, sir, and still more emphatically to
ourselves and to our nation (of the language of whose hearts we
presume to think ourselves at this moment the faithful interpreters)
to express the sentiments with which it is contemplated.
The spectacle of a whole nation, the freest and most enlightened in
the world, offering by its Representatives the tribute of unfeigned
approbation to its first citizen, however novel and interesting it may
be, derives all its lustre—a lustre which accident or enthusiasm could
not bestow, and which adulation would tarnish—from the
transcendent merit of which it is the voluntary testimony.
May you long enjoy that liberty which is so dear to you, and to which
your name will ever be so dear. May your own virtues and a nation's
prayers obtain the happiest sunshine for the decline of your days
and the choicest of future blessings. For your country's sake—for the
sake of Republican liberty—it is our earnest wish that your example
may be the guide of your successors; and thus, after being the
ornament and safeguard of the present age, become the patrimony
of our descendants.
Mr. Venable observed, on a paragraph wherein it speaks of the
"tranquillity of the interior frontier," he did not know what was the
meaning of the expression: he moved to insert "Western frontier" in
its stead.
Mr. Ames observed that the words of the report are in the President's
Speech; however, he thought the amendment a good one. It then
passed.
In the fourth paragraph are these words: "Your endeavors to fulfil
that wish cannot fail, therefore, to interest our attention." At the
word "wish," Mr. Giles proposed to insert these words: "and by all
honorable means to preserve peace, and restore that harmony and
affection which have heretofore so happily subsisted between the
French Republic and this country;" and strike out the words that
follow "wish" in that paragraph. He said, his reasons for moving this
amendment were to avoid its consequences. He really wished the
report entirely recommitted, as there were many objectionable parts
in it. He had been very seriously impressed with the consequences
that would result from a war with the French Republic. When I
reflect, said Mr. G., on the calamities of war in general, I shudder at
the thought; but, to conceive of the danger of a French war in
particular, it cuts me still closer. When I think what many gentlemen
in mercantile situations now feel, and the dreadful stop put to
commerce, I feel the most sincere desire to cultivate harmony and
good understanding. I see redoubled motives to show the world that
we are in favor of a preservation of peace and harmony.
Mr. W. Smith said, he should not object to the amendment; but he
thought it only an amplification of a sentiment just before expressed.
He did not see any advantage in the sentiment as dilated, nor could
he see any injury which could accrue from it. He hoped every
gentleman in the House wished as sincerely for the preservation of
peace as that gentleman did.
Mr. Ames wished to know of the gentleman from Virginia, whether he
meant to strike out the latter part of this paragraph; if he did, he
would object to it.
Mr. Giles said, he did not mean to strike out any more of this
paragraph.
Mr. Ames wished it not to be struck out. By the amendment to strike
out, we show the dependence we place on the power and protection
of the French. While we declare ourselves weak by the act, we lose
the recourse to our own patriotism, and fly, acknowledging an
offence never committed, to the French for peace. He hoped the
gentleman would be candid upon this occasion.
Mr. Giles said, he only wished this House to express their most
sincere and unequivocal desire in favor of peace, and not merely to
leave it to the President. He said, he had spoken upon this occasion
as he always had done on this floor. He always had, and he hoped
always should state his opinions upon every subject with plainness
and candor.
The amendment passed unanimously.
Mr. Giles then proposed an amendment to the latter part of the same
paragraph which would make it read thus: "We cherish the pleasing
hope that a spirit of mutual justice and moderation will ensure the
success of your perseverance." The amendment was to insert the
word "mutual." He thought we ought to display a spirit of justice and
moderation as well as the French. This amendment, he thought,
would soften the expression, and, acting with that spirit of justice
and moderation, accomplish a reconciliation. The amendment was
adopted.
On the Chairman's reading the last paragraph except one in the
report, which reads thus: "The spectacle of a whole nation, the
freest and most enlightened in the world," Mr. Parker moved to strike
out the words in italic. Although, said he, I wish to believe that we
are the freest people, and the most enlightened people in the world,
it is enough that we think ourselves so; it is not becoming in us to
make the declaration to the world; and if we are not so, it is still
worse for us to suppose ourselves what we are not.
Mr. Harper said he had a motion of amendment in his hand which
would supersede the necessity of the last made, which, if in order,
he would propose: it was to insert words more simple. He thought
the more simple, the more agreeable to the public ear. His
amendment, he thought, would add to the elegance and conciseness
of the expression. He did not disapprove of the Address as it now
stood, but he thought it might be amended. This, he said, would add
to the dignity, as well as to the simplicity of the expression. He
thought it would be improper to give too much scope to feeling:
amplitude of expression frequently weakens an idea.
Mr. Giles said he saw many objectionable parts in the amendments
proposed by the gentleman just sat down. He wished to strike out
two paragraphs more than Mr. Harper had proposed; indeed, he
wished the whole to be recommitted, that it might be formed more
congenial to the wishes of the House in general, and not less
agreeable to the person to be addressed.
Mr. Smith observed, that as the answer had been read by paragraphs
nearly to the close, he thought it very much out of order to return to
parts so distant.
The Chairman said that no paragraph on which an amendment had
been made could be returned to; but where no amendment had
been made, it was quite consistent with order to propose any one
gentlemen may think proper.
Mr. W. Smith opposed striking out any paragraph. It was, he said, the
last occasion we should have to address that great man, who had
done so much service to his country. The warmth of expression in
the answer was only an evidence of the gratitude of this House for
his character. When we reflect on the glowing language used at the
time when he accepted of the office of President, and at his re-
election to that office, why, asked he, ought not the language of this
House to be as full of respect and gratitude now as then? particularly
when we consider the addresses now flowing in from all parts of the
country. I object to the manner of gentlemen's amendments as
proposed, to strike out all in a mass. If the sentiments were
agreeable to the minds of the House, why waste our time to alter
mere expressions while the sentiment is preserved? No doubt every
gentleman's manner of expression differed, while their general ideas
might be the same. He hoped mere form of expression would not
cause its recommitment.
Mr. Giles did not object to a respectful and complimentary Address
being sent to the President, yet he thought we ought not to carry our
expressions out of the bounds of moderation; he hoped we should
adhere to truth. He objected to some of the expressions in those
paragraphs, for which reason he moved to have the paragraphs
struck out, in order to be amended by the committee. He wished to
act as respectful to the President as any gentleman, but he observed
many parts of the Address which were objectionable. It is unnatural
and unbecoming in us to exult in our superior happiness, light, or
wisdom. It is not at all necessary that we should exult in our
advantages, and thus reflect on the unhappy situation of nations in
their troubles; it is insulting to them. If we are thus happy it is well
for us; it is necessary that we should enjoy our happiness, but not
boast of it to all the world, and insult their unhappy situation.
As to those parts of the Address which speak of the wisdom and
firmness of the President, he must object to them. On reflection, he
could see a want of wisdom and firmness in the Administration for
the last six years. I may be singular in my ideas, said he, but I
believe our Administration has been neither wise nor firm. I believe,
sir, a want of wisdom and firmness has brought this country into the
present alarming situation. If after such a view of the Administration,
I was to come into this House and show the contrary by a quiet
acquiescence, gentlemen would think me a very inconsistent
character. If we take a view of our foreign relations, we shall see no
reason to exult in the wisdom or firmness of our Administration. He
thought nothing so much as a want of that wisdom and firmness had
brought us to the critical situation in which we now stand.
If it had been the will of gentlemen to have been satisfied with
placing the President in the highest possible point of respect amongst
men, the vote of the House would have been unanimous, but the
proposal of such adulation could never expect success. If we take a
view into our internal situation, and behold the ruined state of public
and private credit, less now than perhaps at any former period
however, he never could recollect it so deranged. If we survey this
city, what a shameful scene it alone exhibits, owing, as he supposed,
to the immense quantity of paper issued. Surely this could afford no
ground for admiration of the Administration that caused it.
I must acknowledge, said Mr. Giles, that I am one of those who do
not think so much of the President as some others do. When the
President retires from his present station, I wish him to enjoy all
possible happiness. I wish him to retire, and that this was the
moment of his retirement. He thought the Government of the United
States could go on very well without him; and he thought he would
enjoy more happiness in his retirement than he possibly could in his
present situation. What calamities would attend the United States,
and how short the duration of its Independence, if one man alone
can be found to fill that capacity! He thought there were thousands
of citizens in the United States able to fill that high office, and he
doubted not that many may be found whose talents would enable
them to fill it with credit and advantage. Although much had been
said, and that by many people, about his intended retirement, yet he
must acknowledge he felt no uncomfortable sensations about it; he
must express his own feelings, he was perfectly easy in prospect of
the event. He wished the President as much happiness as any man.
He declared he did not regret his retreat; he wished him quietly at
his seat at Mount Vernon; he thought he would enjoy more
happiness there than in public life. It will be very extraordinary if
gentlemen, whose names in the yeas and nays are found in
opposition to certain prominent measures of the Administration,
should come forward and approve those measures: this we could not
expect. He retained an opinion he had always seen reason to
support, and no influence under Heaven should prevent him
expressing his established sentiments; and he thought the same
opinions would soon meet general concurrence. He hoped
gentlemen would compliment the President privately, as individuals;
at the same time, he hoped such adulation would never pervade that
House.
I must make some observation, said Mr. G., on the last paragraph
but one, where we call ourselves "the freest and most enlightened
nation in the world:" indeed, the whole of that paragraph is
objectionable; I disapprove the whole of it. If I am free, if I am
happy, if I am enlightened more than others, I wish not to proclaim
it on the house top; if we are free, it is not prudent to declare it; if
enlightened, it is not our duty in this House to trumpet it to the
world; it is no Legislative concern. If gentlemen will examine the
paragraph, [referring to that contained in the parenthesis,] it seems
to prove that the gentleman who drew it up was going into the field
of adulation; which would tarnish a private character. I do think this
kind of affection the President gains nothing from. The many long
Addresses we hear of, add nothing to the lustre of his character. In
the honor we may attempt to give to others we may hurt ourselves.
This may prove a self-destroyer; by relying too much on
administration, we may rely too little on our own strength.
Mr. Sitgreaves said, that whatever division of the question gentlemen
would propose, was indifferent to him; the words of the answer
were perfectly congenial with his wishes, and he was prepared to
give his opposition to any of the amendments proposed. On mature
deliberation, there was not a sentiment in the report but he highly
approved. He could not see any thing unnatural or unbecoming in
drawing just comparisons of our situation with that of our neighbors;
this is the only way we can form a just view of our own happiness. It
is a very necessary way to come to a right knowledge of our own
situation by comparing it with that of other nations. He would not
reproach another people because they are not so happy as we are;
but he thought drawing simple comparisons in the way of the report
was no reproach. He was not against bringing the comparison down
to private life, as the gentleman from Virginia had done; he should
think it wrong in a man to exult over his neighbor who was
distressed or ignorant, because himself was wealthy or wise. Yet he
saw no impropriety in his own family of speaking of their happiness
and advantages, compared with that of others; it would awaken in
them a grateful sense of their superior enjoyments, while it pointed
out the faults and follies of others, only in order that those he had
the care of may learn to avoid them: thus while our happiness is
pointed out, the miseries of nations involved in distress are
delineated to serve as beacons for the United States to steer clear
of. He did not, with the gentleman from Virginia, in any degree,
doubt of the wisdom or firmness of the Administration of America. In
the language of the Address, he entertained a very high opinion of
it, "a grateful conviction that the wise, firm, and patriotic
Administration of the President had been signally conducive to the
success of the present form of Government." Such language as this
is the only reward which can be given by a grateful people for labors
so eminently useful as those of the President had been. This was not
his sentiment merely, it was the sentiment of the people of America.
Every public body were conveying their sentiments of gratitude
throughout the whole extent of the Union. Why then should this
House affect a singularity, when our silence on these points would
only convey reproach instead of respect. If these sentiments were
true, why not express them? But if, on the contrary, what the
gentleman asserted, that the Administration of the President had
been neither wise, firm, nor patriotic, then he would concur with the
motion for striking out; but he was not convinced of the truth of this
assertion; and while this is not proved, he should vote against the
motion.
Mr. Sitgreaves said, he could not agree with the motion of the
gentleman from South Carolina, (Mr. Harper,) because his motion
was for substituting other words in the place of those in the report,
without any reason whatever. If the gentleman, by altering the
phraseology, can make the sentiment any better, by all means let it
be done: but if the sentiment is not to be changed, why alter it
merely to substitute other words? On the whole, Mr. S. observed,
that he did not see the answer could in any degree be reproached.
There are no sentiments in it but what are justifiable on the ground
of truth; they are free from adulation. It is such an expression of
national regret and gratitude as the circumstance calls for; a regret
at the retirement of a faithful and patriotic Chief Magistrate from
office. A regret and gratitude which he believed to be the sentiment
of Americans.
Mr. Swanwick began by observing that there were points in the
Address in which all gentlemen seemed to agree, while on other
parts they cannot agree. We all agree in our desire to pay the
President every possible mark of respect; but we very materially
disagree wherein a comparison is drawn between this and foreign
nations. If we are happy and other nations are not so, it is but well
for us; but he thought it would be much more prudent in us to let
other nations discover it, and not make a boast of it ourselves. It is
very likely that those nations whom we commiserate may think
themselves as happy as we are: they may feel offended to hear of
our comparisons. If we refer to the British Chancellor of the
Exchequer in his speeches, he would tell us that is the happiest and
most prosperous nation upon earth. How then can we commiserate
with it as an unfortunate country? If, again, we look to France, that
country which we have pointed out as full of wretchedness and
distress, yet we hear them boast of their superiority of light and
freedom, and we have reason to believe not without foundation. A
gentleman had talked about the flourishing state of our agriculture,
and asserted that our late commercial calamities were not proofs of
our want of prosperity, which the gentleman compared to specks in
the sun. That gentleman speaks as though he lived at a distance.
Has he heard of no commercial distresses, when violations so
unprecedented have of late occurred? One merchant has to look for
his property at Halifax, another at Bermuda, another at Cape
Françoise, another at Gonaives, &c.; all agree that they have
suffered, and that by the war. These are distresses gentlemen would
not like to feel themselves. Mr. S. said he had felt for these
occurrences. We are not exempt from troubles: probably we may
have suffered as much as other nations who are involved in the war.
It is a question whether France has been distressed at all by the war.
She has collected gold and silver in immense quantities by her
conquests, together with the most valuable stores of the productions
of the arts; as statues, paintings, and manuscripts of inestimable
worth; and at sea has taken far more in value than she has lost:
besides, her armies are subsisting on the requisitions her victories
obtain. And has England gained nothing by the war? If we hearken
to Mr. Pitt, we may believe they are very great gainers. Surely the
islands in the West and East Indies, Ceylon, and the Cape of Good
Hope, the key to the East Indies, are advantages gained; besides
the quantity of shipping taken from our merchants. Mr. S. thought if
we were to compare, we should find those nations had gained by
the war, while we had lost; and of course there was no reason for us
to boast of our advantages.
Mr. W. Smith next rose, and observed that gentlemen wished to
compliment the President, but took away every point on which
encomium could be grounded. One denies the prosperity of the
country, another the free and enlightened state of the country, and
another refuses the President the epithet of wise and patriotic.
Mr. Giles here rose to explain. If he was meant, he must think the
gentleman was wrong in his application. He said he had never
harbored a suspicion of the good intentions of the President, nor did
he deny his patriotism; but the wisdom and firmness of his
Administration he had doubted. He thought him a good meaning
man, but often misled.
Mr. Smith again rose, and said, he must confess himself at a loss for
that refinement to discover between the wisdom and patriotism of
the President, and that of his Administration. It was moved to strike
out this acknowledgment of wisdom and firmness. What were we to
substitute as complimentary to him in its place? The first paragraph
proposed to be struck out related to our speaking of the tranquillity
of this country, compared to nations involved in war. Could this give
offence, because we feel pleasure in being at peace? It was only
congratulating our own constituents on the happiness we enjoy. To
appreciate the value of peace, it was necessary to compare it with a
state of war. It was the wisdom of this country to keep from war,
and other nations hold it up as exemplary in us. The gentleman
himself has declared his wish for the preservation of peace; and
though he admires it, and nations admire it in us, yet we are not to
compare our state with nations involved in the calamities of war, in
order to estimate our enjoyments. The words of this Address are not
a communication to a foreign minister, it is a congratulation to our
own Chief Magistrate of the blessings he, in common with us,
enjoys. Mr. S. hoped the words would not be struck out.
Mr. Dayton (the Speaker), said, that he did not rise to accept the
challenge given by the gentleman who spoke last from South
Carolina, and to point out a nation more free and enlightened than
ours; nor did he mean to contest the fact of ours being the freest
and most enlightened in the world, as declared in the reported
Address, but he was nevertheless of opinion that it did not become
them to make that declaration, and thus to extol themselves by a
comparison with, and at the expense of all others. Although those
words were in his view objectionable, he was far from assenting to
the motion for striking out the seven or eight last clauses of the
Address. The question of order having been decided, Mr. D. said he
would remind the committee, that if they wished to retain, or even
to amend, any section or sentence of all that was proposed to be
struck out, they ought to give their negatives to this motion, as the
only means of accomplishing their purpose. It was sufficient,
therefore, for those who were opposed to the question for striking
out the whole, to show that any part included within it ought to be
preserved. Not unnecessarily to waste time, by lengthening the
debate, he would take the clause first in order, and confine his
remarks to that alone. This part of the Address had certainly not
been read, or had been misunderstood and misrepresented by the
member from Pennsylvania.
Mr. Ames said, if gentlemen meant to agree to strike out the whole as
proposed, in order to adopt those words substituted by the
gentleman from South Carolina (Mr. Harper), he must observe that
he thought this would be as far from giving satisfaction to others,
who, it appeared, wanted no substitute. He, therefore, hoped that
kind of influence would not prevail on this occasion. The gentleman
who made the motion did it to accommodate matters, and not
because he himself objected to the answer reported.
It is well known that a committee of five members, opposite in
sentiment, was appointed to prepare a respectful Address in answer
to the President's Speech. [Here the original instructions were read.]
As it was the duty of the committee to prepare a respectful Address,
it cannot be matter of surprise, although it may of disapprobation
with some, that the committee did their duty, and have taken notice
of the several matters recommended to the House in that Speech.
Respecting the particular notice they have taken, it might have been
thought that some difficulty would occur. He said he need not
observe, that the committee had reason to imagine that the form of
the report would be agreeable to the House, as they were
unanimous; although there had been in the wording some little
difference of opinion, yet all agreed substantially in the Address,
from a conviction of the delicacy of the subject. For that reason, if
that only, unless the sentiments in the report of the Address should
be found inconsistent with truth, he hoped no substitute of a form of
words merely would prevail, as it would no longer be that agreed to
in the committee, nor could come under their consideration equal to
the printed report. He therefore trusted that when the committee
came to the question, whether to strike out or not, gentlemen would
be guided by no other motive to vote for striking out, than an
impropriety in the sentiments through an evident want of truth in
them; and if such cannot be discovered, why strike out the
expressions?
It had been observed by some gentleman, that the cry of foreign
influence is in the country. He did not see such a thing exist. He
would not be rudely explicit as to the foundation there was for such
a cry; but when it was once raised, the people would judge whether
it was fact or not. He could not tell how this influence was produced,
but the world would draw a view how far we were under foreign
influence. Mr. A. here alluded to the influence which foreign agents
wished to have over the minds of the people of this country, in order
to support a factious spirit, probably to the appeal lately made to the
people. He also alluded to a circumstance when the Imperial Envoy,
M. Palm, in 1727, at London, published a rescript, complaining of the
conduct of that Court; the spirit of the nation rose, and discord was
sown. In consequence of which the Parliament petitioned the King to
send the Envoy out of the country for meddling with the concerns of
their nation. That is the nation which we call corrupted. Yet a similar
affair has occurred here, and it is not to be reprobated; we are not
to complain of it, nor even hear it, according to this doctrine.
Independence is afraid of injuries, and almost of insults. We must
forbear to exult in our peace, our light, our freedom, lest we should
give offence to other nations who are not so. This may be the high
tone of independence in the views of some people, but I must
confess it is not so in mine; but it is probable those people may be
wiser than I am, and their views extend farther. Foreign influence
exists, and is disgraceful indeed, when we dare admire our own
constitution, nor adore God for giving us to feel its happy effects. He
thought, respecting the recent complaints of the French Minister,
that there was not even a pretext for the accusation.
It had been observed by a gentleman, that the President, no doubt,
is a very honest man, and a patriot, but he did not think him a wise
man.
Mr. Giles here rose to explain. He said that, in his assertions, he
meant not to reflect on his private character. He referred to his
Administration. No doubt but the gentleman possessed both.
Mr. Ames said, he considered well what the gentleman had said. As a
private man, his integrity and goodness cannot be doubted; but in
his Administration—here we are to stop short; not a word about
that; it won't bear looking into; it has been neither firm nor wise. If
the House, in their Address to him, were to say, we think you a very
honest, well-designing man, but you have been led astray,
sometimes to act treacherously, and even dishonest in your
Administration—we think you a peaceful man, and though much
iniquity may have been practised in your Government, yet we think
you are not in fault; on the whole, sir, we wish you snugly in
Virginia. Such sentiments as these I do not like. Is this an Address or
an insult? Is this the mark of respect we ought to show to the first
man in the nation? Mr. A. observed, that he did not agree with the
gentleman from South Carolina (Mr. Smith), who said, that the
President would carry daggers in his heart with him into his retreat
from public life, if we refuse him our testimony of gratitude. No, he
bears in his breast a testimony of his purity of motive; a conscious
rectitude, while in public life, which daggers could not pierce. He
would retire with a good conscience; perhaps it would be said this
was adulation, but let it be remembered this was truth; this was not
flattery; let gentlemen deny this; let them prove that this is not the
will of their constituents. The country would judge our opinions
when we come to give our yeas or nays; then the real friends of that
man would be known.
The gentleman wishes him back to Virginia, was glad he designed to
go; he did not regret his resignation. His name will appear in that
opinion. The whole of the President's life would stamp his character.
His country, and the admiring world knew it; and history keeps his
fame, and will continue to keep it. We may be singular in our
opinions of him, but that will not make his character with the world
the less illustrious. We now are to accept of his resignation without a
tribute of respect. We are not to speak of him as either wise or firm.
We can only say he is an honest man: this would scarcely be
singular; many a man is honest without any other good
qualifications. What circle would gentlemen fix the committee in to
amend this Address, if they are not to give scope to these
sentiments? Better appoint no committee at all. If we address the
President at all, I hope it will be respectfully, for loth respect is insult
in disguise. I hope we shall not alter the original draft of the
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