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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: peter.szoevenyi@uzh.ch

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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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
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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
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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
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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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The Project Gutenberg eBook of An index to
the collected works of William Hazlitt
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Title: An index to the collected works of William Hazlitt

Author: William Hazlitt

Editor: Arnold Glover

A. R. Waller

Release date: November 26, 2023 [eBook #72232]

Language: English

Original publication: London: J. M. Dent & Co, 1906

Credits: Richard Tonsing and the Online Distributed

Proofreading Team at https://www.pgdp.net (This file
was produced from images generously made available
by The Internet Archive)

*** START OF THE PROJECT GUTENBERG EBOOK AN INDEX

TO THE COLLECTED WORKS OF WILLIAM HAZLITT ***
Transcriber’s Note:
New original cover art included with this eBook is
granted to the public domain.
INDEX TO HAZLITT’S
COLLECTED WORKS

All Rights Reserved

AN INDEX TO THE COLLECTED WORKS OF

WILLIAM HAZLITT

EDITED BY A. R. WALLER
AND ARNOLD GLOVER
❦

1906
LONDON: J. M. DENT & CO.
29 AND 30 BEDFORD STREET, W.C.
 TO THE MEMORY OF
A. G.

E. H. G.
A. R. W.
 PREFACE

It was originally intended to add an index at the end of volume xii.

of our edition, but so many of Hazlitt’s previously unidentified
writings were discovered during the progress of the work, that it was
found to be impossible to make volumes xi. and xii. any thicker. A
brief index of the titles of Hazlitt’s various essays and papers closed,
therefore, volume xii., and the publication of a separate index was
designed.
The scope of this index was agreed upon between Arnold Glover
and myself; the slips were made under my direction, and it was my
intention to have checked them myself. Upon the death of Arnold
Glover, however, I had to undertake other work left uncompleted by
him, and the labour of checking each entry in the proofs with the
twelve volumes indexed has been accomplished by Mrs Arnold
Glover.
The book is now published, through the collaboration of Mrs
Arnold Glover, Messrs J. M. Dent & Co. and myself, at a price
considerably below cost, in order to place it within reach of all lovers
of Hazlitt; and in memory of one who spared no pains in his self-
chosen task of making the writings of Hazlitt better known, one who
added so considerably to the known bulk of those writings.
Recognition of the value of his labour has, since his death, been
ungrudgingly given him. It remains to be added that he himself
would have regarded that labour as incomplete until he had passed
the last proof-sheet of this index.

A. R. Waller.

Cambridge, 1st September 1906.

 NOTE

The index is practically confined to the names of persons and

things. The names of imaginary persons, characters in novels, plays,
etc., are indexed under the first word of the name by which they are
usually designated, e.g. Tony Lumpkin, Don Quixote, Dr Faustus.
The names of real persons are indexed in the ordinary way. In the
very heavy list of Quotations the method adopted has followed so far
as seemed possible that in use in the Index to Quotations in Notes
and Queries. The list of quotations has been very considerably
lightened by leaving out almost all phrases from Shakespeare and
Milton: to have added these would have made the list of quotations
almost double its present length, so often did Hazlitt quote from
these two. Hazlitt’s spelling has been followed where deemed
characteristic.
It has been discovered that a short essay has, by mistake, been
printed twice in the text (vol. vii. p. 360, and vol. xi. p. 433). Owners
of the edition will, perhaps, be so good as to note this fact in their
copies.
It is the intention of the surviving editor to keep the notes and
index up to date, in case a second impression should be required. He
will be obliged if readers will send him any suggestions they may
wish to make.
 ADDENDA

Beyle, M. H., vi. 285; viii. 411.

For Sir Thomas Booby, see under
Sir Thomas, not Booby.
Bothwell, v. 142.
Change-Alley, iii. 297.
Conciones ad Populum, x. 131.
Destut, etc., read Destutt de Tracy,
and add vii. 323.
Dr Slop, viii. 121.
Ecole des Femmes, viii. 554; xi. 276.
Excursion, The, vii. 76.
French Revolution (Burke’s), ix.
473 n.; xii. 291.
For Still substitute Stevenson as the probable author of Gammer
Gurton’s Needle.
Geneva, ix. 216.
Junius, iv. 217.
For La Fontaine the fabulist, see
Under La, not Fontaine.
Madge Wildfire, viii. 413 n.
Neapolitan Nobleman, xi. 222.
Onslow, Arthur, vii. 271.
Present Discontents, iii. 335.
For Sterne’s Uncle Toby, See Under
“My Uncle,” as well as Uncle
Toby, and add v. 129.
Stoddart, Dr, xi. 444.
 ADDENDA TO QUOTATIONS
above all pain, all passion, and all pride, ix. 59.
all this world were one glorious lie, v. 334.
and doubtless ’mong the grave and good, vii. 366.
as good as a prologue, viii. 309.

calm pleasures, vii. 318.

commanded to shew the knight in love, i. 348.
constrained by mastery, i. 151.

deem not devoid of elegance, vii. 317.

each other’s beams to share, xi. 488.

earth destroys those raptures, vii. 318.
elegant Petruchio, etc., i. 344.
Elysian beauty, vii. 320.
endure having hot molten lead, etc., vii. 322.

first garden ... innocence, i. 105.

for a song, vii. 362.

gentlemen’s gentlemen, vii. 211.

glared round his soul, vii. 319.
grand carnival of this our age, xi. 440.
Hamlet, to leave the part of, xii. 383.
he was hurt and knew it not, vii. 354.
head to the East, vii. 342.
his face ’twixt tears and smiles, xi. 480.
his grace looks cheerfully, viii. 183.
his look made the still air cold, vii. 99.
huge, dumb heap, viii. 448.

interlocutions between Lucius and Caius, iv. 276.

is it to be supposed that it is England, xi. 444.

leave all and follow it, vii. 315.

license of the time, i. 235.
like dew-drops from the lion’s mane, v. 267 n.
like poppies spread, vii. 308.

madness, that fine, xii. 340.

meek mouths ruminant, iii. 239.
mighty dead, vii. 365.
mind reflecting ages past, iv. 213.
mouth with slumbery pout, viii. 478.

No maid could live near such a man, i. 305.

No, thou art not my child, viii. 427.
Not Fate itself could awe, xi. 410.
now of the planetary, iv. 230.
Oh, not from you, vii. 339.
out on the craft, vii. 365 n.
owes no allegiance, i. 112.

paint a sunbeam to the blind, etc., v. 237.

perceive a softness, etc., xi. 522.
picks pears, etc., ix. 71.
play at bowls with the sun and moon, ix. 64.
play with wisdom, xi. 551.
pomp of elder days, v. 177.
prevailing gentle arts, iii. 108.
proper study, etc., vii. 312.

rejoice when good kings bleed, i. 191.

right divine, iii. 288.
roast me these Violantes, viii. 156.
round which with tendrils, etc., vii. 310.

sailing with supreme dominion, vii. 339.

see o’er the stage, etc., vi. 273; xii. 123.
spin his brains, vii. 319.
stand all apart, viii. 181.
still, small, etc., vii. 336.
strange that such difference, etc., xi. 505.
sweet passion of love, viii. 261.

there were two upon the housetops, viii. 393.

thin partitions do their bounds, etc., viii. 217.
trees in Sherwood forest, v. 143.

well of English, etc., vii. 321.

what was my pride, etc., xi. 455.
which waste the marrow, xii. 427.
whose body nature was, vii. 320.
winged words, xii. 345.
within our bosoms, etc., ii. 395.

you ask her crime, vii. 350.

About one-third of the above are additional occurrences of

quotations already indexed.
 A.

A. (Mr), vi. 390.

Aaron (Shakespeare’s Titus Andronicus), i. 357; v. 207.
Aaron’s Rod, iv. 63.
Abbot’s Ford, iv. 245.
Abbott, William, vi. 277; viii. 235, 266, 292, 333, 335, 413, 455; xi.
402.
—— Mr Speaker (Lord Colchester), iv. 199; xii. 370.
À Becket, Thomas, v. 143.
A’Beckett, Gilbert, v. 143.
Abednego, iii. 265.
Abel Drugger (Ben Jonson’s The Alchemyst), ii. 76 n., 79; vi. 273,
418; viii. 227, 228; xii. 33.
Abelard and Eloise (Pope’s), v. 75.
—— Peter, iv. 213; v. 75; vii. 96; ix. 146, 354; xii. 165.
Aben Hamet (in Dimond’s Conquest of Taranto), viii. 368.
Aberdeen, ii. 209; viii. 290; ix. 492.
Abernethy, John, vi. 62.
Abhorson (in Shakespeare’s Measure for Measure), i. 346.
Abimelech Henley (in Holcroft’s Anna St Ives), ii. 128.
Abinger, Lord (Mr Scarlett), vii. 174 n., 516.
Abington, Mrs, i. 157; vi. 426, 453; viii. 74, 174, 393; ix. 147; xii. 24.
Aboam (in Southerne’s Oroonoko), xi. 302.
Abraham Adams (in Fielding’s Joseph Andrews), viii. 106; x. 328.
Abruzzi, The Passes of the, ix 255, 259; x. 283, 291.
Absalom and Achitophel (Dryden), v. 80.
Abstract Ideas, On, xi. p. v., 1.
—— —— xi. 1.
Abstraction, On, xi. 180.
Academie de Musique, viii. 363.
Academy, at Bologna, ix. 205, 264.
—— of Compliments, i. 235.
—— of Painting (Venice), ix. 273, 274.
—— of St Luke, x. 280, 296.
Accusation. See Anglade Family.
Acetto, Count, ix. 213.
Achilles, i. 221; iv. 225; v. 15, 54, 64; vii. 255; x. 6, 98; xii. 8.
Achitophel (Dryden’s), iii. 400; v. 80.
Achmet (in John Brown’s Barbarossa), viii. 372.
Acis and Galatea (by John Gay), vii. 103.
Ackerman, Rudolf, i. 366; viii. 143; ix. 313.
Acropolis, ix. 379; x. 343; xi. 227, 486.
Actæon—Hunting (Titian), i. 27, 78.
Acted Drama in London, Essays on the, viii. 381.
Actium, i. 229; v. 50; viii. 192.
Acton, Sir Francis, v. 213.
Actor, The (Holcroft’s paper), ii. 87.
Actors and Acting, On, i. 153, 156, 382.
—— ought to sit in the Boxes? Whether, vi. 272.
Actors and the Public, xi. 348.
Acts of Uniformity, xi. 314.
Adair, Sir R., ii. 169, 214, 227.
Adam, i. 38, 105 seq., 385, 425; iv. 337; v. 60, 66; vi. 96, 396, 411; vii.
36; xi. 233, 452; xii. 276, 455 n.
—— (in Shakespeare’s As You Like It), i. 340.
—— (Michael Angelo’s), ix. 241, 332, 362.
—— (in Milton’s Paradise Lost), v. 357, 371.
—— and Eve (Barry’s), ix. 418.
—— —— (Martin’s), vi. 398; vii. 292 n.; xi. 553.
—— —— in Paradise (Poussin), vii. 291, 292; ix. 109.
—— —— driven out of Paradise (Ribera’s), ix. 70.
Adam Bell (in Holcroft’s The Noble Peasant), ii. 110.
Adams, Mr (Sec. of the Constitutional Soc.), ii. 153.
Addington, Henry, Viscount Sidmouth, iii. 3, 38; viii. 17; xii. 206,
450.
—— Hiley, iii. 238, 300.
Addison, Joseph, i. 2, 8, 9, 370, 372, 374, 380, 382, 415; ii. 43, 212;
iv. 233, 269, 367; v. 78, 319, 373; vii. 6, 36; viii. 22, 89, 97, 98, 99,
101, 105, 158; ix. 276; x. 112, 206, 359; xi. 391, 489, 543, 546; xii.
35, 170, 330, 331, 375.
Address to the Inhabitants of New Lanark (Robert Owen’s), iii. 121.
Adela (in Holcroft’s The Noble Peasant), ii. 110.
Adelaide, or The Emigrants (Sheil’s), viii. 308, 530.
Adelard (in L. Bonaparte’s Charlemagne), xi. 232.
Adelinde (in L. Bonaparte’s Charlemagne), xi. 235.
Adelphi, i. 35; viii. 404, 411, 412; ix. 420, 421, 422; x. 200; xi. 242;
xii. 168, 195, 196.
Adige, The, ix. 277.
Admetus, iii. 206.
Admirable Crichton, viii. 459; xii. 34.
Admiralty, vii. 115.
Adonis, Gardens of, v. 38.
Adoration, An (Raphael’s), ix. 73.
—— of the Angels (Poussin’s), ix. 51.
—— of the Shepherds (Caracci’s), ix. 26.
Adrian (Pope), xi. 235.
Adriana (in Shakespeare’s Comedy of Errors), i. 352.
Adriano de Armada (in Shakespeare’s Love’s Labour’s Lost), v. 324.
Adrian’s Villa, ix. 256; xi. 199.
Adriatic, ix. 200, 264, 266, 284.
Adventurer, The (Hawksworth’s), vii. 226; viii. 104.
Adventures of Friar Albert (Chaucer), xii. 30.
—— at Madrid, or Gallantry (a play), viii. 399.
Advertisement and Biographical and Critical Notes from the
Eloquence of the British Senate, iii. 388.
Advice to a Patriot; in a Letter addressed to a Member of the Old
Opposition, iii. 1.
Ægeon of Syracuse (Shakespeare’s Comedy of Errors), iv. 341; vi. 58.
Æmilia (in Two Noble Kinsmen), v. 261.
—— (Shakespeare’s Othello), i. 201 seq., 293; vi. 303.
Æneas, i. 416; iii. 461; v. 54; ix. 171; x. 20; xi. 492; xii. 19.
—— Escaping, etc. (Barry’s), ix. 415.
—— (in Opera Didona Abandonnata), viii. 197.
—— and Dido (Guérin’s), ix. 135.
Æneid (Gawin Douglas’ version), v. 399.