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A PROJECT REPORT ON

“study of diversity of trichomes”

SUBMITTED

TO

DEPARTMENT OF BOTANY
RAJARSHI SHAHU MAHAVIDYALAYA, LATUR
(AUTONOMOUS)

FOR THE AWARD OF

B.Sc1st year
BY

Mr.Gaikwad Ganesh (B.Sc. I)

UNDER THE GUIDANCE OF

Mr.D Awad
DEPARTMENT OF BOTANY

RAJARSHI SHAHU MAHAVIDYALAYA, LATUR

(AUTONOMOUS)

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RAJARSHI SHAHU MAHAVIDYALAYA, LATUR.

(AUTONOMOUS)
DEPARTMENT OF BOTANY

DECLARATION

I hereby declare that the project entitled “study of diversity of

trichomes” submitted to Department of Botany, Rajarshi Shahu
Mahavidyalaya (Autonomous), Latur during 2024-2025 for the award
of B.Sc. I Year degree is written by me and has not previously been
formed on the basis for the award of any degree or diploma or other
similar title of this or any other University, or examining body.

Place: Latur Mr. Gaikwad Ganesh

B.Sc..I

Project Student

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CERTIFICATE
This is to certify that the project report entitle Effect of NPK on Monocot and Dicot

Plant which is being submitted here with is the result of the project work completed

by Mr.Gaikwad Ganesh Sandipan under my supervision and guidance and to the

best of our knowledge and belief the work embodied in the report has not formed

earlier the basis for the award of any other university or examining body.

Place : Latur

Head Project Guide

Prof. S. N. Shinde Mr.A.Dnyanoba

Department of Botany Department of Botany
Rajarshi Shahu Mahavidyalaya, Latur. Rajarshi Shahu Mahavidyalaya, Latur.

Dr. M. H. Gavhane
Principal
Rajarshi Shahu Mahavidyalaya (Autonomous) Latur.

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ACKNOWLEDGEMENTS

I would like to thank the many people who helped me to complete this project.
My advisor, Mr.Awad Dnyanoba Department of Botany, Rajarshi Shahu
Mahavidyalaya, Latur for her valuable guidance to complete this project. I am
thankful to Prof. S.N. Shinde, Head, Department of Botany for always being
available to help me.
I sincerely thank Dr. M.H. Gavhane, Principal, Rajarshi Shahu
Mahavidyalaya, Latur, for his kind support and making the computer lab, internet,
library facilities available to me. My deep appreciation to Prof. S. N. Shinde Vice
Principal, Rajarshi Shahu Mahavidyalaya, Latur. I own my sincere thanks to most
affectionate and respectful, Dr. K.D. Savant, Mr. D.R. Awad, and Nonteaching staff
of R.S.M.L. Department of Botany, for their kind help and advise for their advice
throughout these years. They are an invaluable source of inspiration and it is their
harmonious nature to be acknowledged. Thanks are due to all my friends studying in
B.Sc. I of Rajarshi Shahu Mahavidyalaya, Latur, for their help.
I am at last of the words while expressing my deep feelings of gratitude
towards my family members for the loving support and inspiration which has
sustained me throughout my life, without their support and encouragement.

Place: Latur

MR. Gaikwad Ganesh B.Sc. I

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INDEX

SR. NO CONTENT PAGE NO.

01 INTRODUCTION

02 REVIEW

03 MATERIAL & METHODS

04 RESULT

05 DISCUSSION &CONCLUSION

06 REFERANCE

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INTRODUCTION
Trichomes, the hairlike protuberances in plants, have been well known to act as the first line of
defense against herbivores, and abiotic stresses, along with other structural defenses such as spines,
thorns, and waxes. We previously reported the tremendous variation in trichome traits among
different wild and cultivated Solanum species and demonstrated that trichomes types and density are
traditionally miscalculated and often misnamed. However, intraspecific variation in trichome traits is
poorly understood, although this has implications for stress tolerance and resistance breeding
programs in economically important crop species and can also mediate ecological interactions at
multiple trophic levels in their wild congeners.
Most of the plants possess hair-like structures that are typically found both above and below the
ground. When these hair-like structures are present on the aerial parts of a plant, they are referred to
as trichomes, a term derived from the Greek word "trichomes" meaning hair. Trichomes can be
divided into two main categories based on their
metabolic activity: glandular trichomes (GTs) and non-glandular trichomes (non-GTs). GTs and non-
GTs are commonly found together on stems, leaves, flowers, and bracts (Rautio et al., 2002).
Glandular trichomes are usually multicellular but unicellular glandular trichomes have also been
reported and they are responsible for synthesizing and storing specialized metabolites such as
polysaccharides, organic acids, proteins, terpenoids, alkaloids, and polyphenols (Schuurink and
Tissier, 2020, Szymanski et al., 2000). They consist of three main parts: the base, stalk, and gland
(Lange, 2015). The gland secretes the metabolites, the stalk contains specialized glands, and the base
connects the stalk to neighboring epidermal cells.
The variation in cell shape and number accounts for the diverse morphological features and sizes
of glandular trichomes. For example, the glandular trichomes of Artemisia annua have a biseriate
shape and are approximately 40– 50 µm long, while tomato glandular trichomes possess long stalks
and measure 2–3 mm in length (Glas et al., 2012, Tissier, 2012). Glandular trichomes can be further
classified as capitate or peltate. Capitate trichomes have an elongated stalk that comprises more than
half of the head height. They exhibit greater variability in stalk length, head morphology, and
secretion patterns compared to peltate trichomes. Peltate trichomes, on the other hand, are shorter in
length and have larger heads composed of 4–18 cells. They possess either unicellular or bicellular
7
stalks (Glas et al., 2012).
Non-glandular trichomes are hair-like structures that protrude from the surface of above-ground
plant organs. Non- glandular trichomes in Arabidopsis include leaf trichomes, stem trichomes, flower
trichomes, fruit trichomes, and hypocotyl trichomes. Differences in size, shape, and density of these
trichomes can be observed by comparing
various Arabidopsis ecotypes and mutant lines. The study of these trichomes in Arabidopsis provides
valuable insights into the development of plants, their defense mechanisms, and their ability to adapt
to their environment (Marks et al., 2009).

The production and distribution of trichomes are regulated either spatially or temporally. In the
early vegetative stages, trichomes are primarily located on the upper surface (adaxial) of the rosette
leaves. However, during the adult

vegetative phase, trichomes can be found on both the upper and lower surfaces (adaxial and abaxial).
As the plant enters the reproductive stage, the number of trichomes decreases on the main
inflorescence stem and flowers (Xiao et al., 2017). Trichome development occurs from precursor cells
in the outermost layer (protodermal) of developing leaf primordia. During this process, neighboring
epidermal cells continue to divide while the initial trichomes separate from the epidermis (Fei et al.,
2020, Han et al., 2022, Mirnezami et al., 2020). Once trichomes exit the mitotic cycle, they undergo
endoreduplication, a phase where DNA replication occurs without nuclear or cellular divisions.
Throughout trichome development, four rounds of endoreduplication take place, resulting in mature
trichomes with approximately three branches and a DNA content of 32 C (1 C representing the DNA
content of an unreplicated haploid genome). During the first round of endoreduplication, trichome
cells change their polarity and begin to protrude from the surface. Concurrently, a second
endoreduplication cycle occurs during the initial branching event. The third round of
endoreduplication takes place after the initiation of the second branching event. Lastly, the fourth
round of endoreduplication corresponds to the expansion of trichome cells, ultimately leading to the
formation of mature trichomes with three branches (Hülskamp et al., 1994). The number of glandular
trichomes on plant leaves can vary based on plant species and their surrounding environmental
factors. In certain cases, when the trichomes are fully developed, their numbers on the leaf are
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relatively fixed, however, in other cases, trichome numbers can be decreased during their expansion
and development in response to external environmental conditions (light, temperature, and humidity),
hormonal factors (gibberellins, cytokinins, and jasmonic acid), and stress responses. These specific
factors and mechanisms can vary among various plant species (Khan et al., 2021).
Some plants possess abundant trichomes on their leaf surfaces, which contribute to the thickening
of the boundary layer between the plant and the environment. This, in turn, helps to reduce heat and
water loss. Trichomes serve as a protective barrier between the plant's epidermal tissues and the
external environment. They absorb harmful UV radiation and release various chemicals that attract
pollinating insects. Non-glandular trichomes act as physical barriers, creating unfavorable
microenvironments for herbivores, parasites, and pathogens. For example, they can form thorn-like
structures to ensnare insects (Riddick and Wu, 2011). Glandular trichomes provide chemical defense
by releasing compounds such as flavonoids, terpenoids, alkaloids, methyl ketones, and surface
proteins (Glas et al., 2012). In response to herbivore attacks, the length and density of trichomes
increase, offering enhanced protection against further damage. Arabidopsis plants experiencing
herbivore attacks can exhibit trichome densities up to 500%, while plants without such attacks
typically have trichome densities of around 20–40% (Dalin et al., 2008). Trichomes act as physical
barriers against various insects by secreting alkaloids that reduce insect populations either by killing
them or inhibiting their growth. Chewing insects are particularly affected by trichomes, as they first
consume the trichomes to access the epidermal layer of the leaf. Consequently, a large accumulation
of trichomes in their digestive systems can lead to their demise (Handley et al., 2005). Trichomes also
play a beneficial role in mitigating drought stress. By increasing the density of trichomes on leaf
surfaces, plants can minimize the harmful effects of drought. Furthermore, trichome density helps to
reduce water transpiration rates (Xiao et al., 2017).

Trichomes play a crucial role in the production of beneficial chemical compounds that are significant
for plant growth and development, disease treatment, and human applications (Kabir et al., 2023,
Qanmber et al., 2019). An excellent example is the presence of trichomes, known as cotton fibers, on
the seed coat of cotton plants (Qanmber et al., 2019, Yang et al., 2020). These fibers are rich in
cellulose, serving as a vital raw material for the textile industry. Similarly, trichomes in plants like
Humulus lupulus, Mentha spp., and A. annua synthesize valuable chemicals that can be utilized in the
extraction of medicines, food additives, and herbicides (Singh et al., 2016). Trichome development is
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controlled by distinct regulatory networks comprising key genes. These conserved networks are
responsible for the intricate process of trichome formation (Pattanaik et al., 2014). Trichome formation
in tobacco (Nicotiana tabacum), is a well-studied process, providing valuable information regarding
the development and function of trichomes in plants. Glandular trichomes in tobacco are important for
their ability to produce and store nicotine, which is a natural insecticide and preventive to herbivores.
Trichomes are the sites for nicotine biosynthesis and storage and the stored nicotine plays a pivotal
role in the defense mechanisms of plants (Howe and Jander, 2008).
The hairs may be subdivided into:
(I) Unicellular; and
(ii) Multicellular.
The unicellular hairs may be un-branched or branched. Multicellular hairs may consist of a single
row of cells or several layers. Some multicellular hairs are branched in dendroid (tree-like) manner;
others have branches oriented largely in one plane (stellate hairs.)
Types of Trichomes:
Stinging Hairs:
They are one of the most interesting types of the trichomes. It contains a poisonous liquid and
consists of a basal bulb like portion from which a stiff, slender and tapering structure is given out.
This tapering structure ends in a small knob like or a sharp point.
The tip is usually somewhat oblique, and as the body of an animal or human being comes in its
contact with some force, the tip is broken off, and the sharp pointed end readily penetrates the skin of
the animal, and fluid is being transferred from the basal knob of the hair to the body of the animal.
Glandular Hairs:
Many plants possess glandular hairs. These hairs may secrete oil, resin or mucilage. A typical
glandular hair possesses a stalk and an enlarged terminal portion, which may be referred to as gland.
The glandular hairs may be uni- or multicellular.
Active secretory cells of glandular trichomes have dense protoplasts and elaborate various
substances, such as volatile oils, resins and mucilage’s, and gums. These substances are excreted and
accumulate between the walls and cuticle. Their final removal from the hair occurs by rupture of the
cuticle.
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Scale or Peltate Hair:

A common type of trichome is the scale, also called peltate hair (from the latin peltatus, target-
shaped or shield like, and attached by its lower surface). A scale consists of a discoid plate of cells,
often borne on a stalk or attached directly to the foot.
Cell Wall of Trichomes:
The cell walls of trichomes are commonly of cellulose and are covered with a cuticle. They may be
lignified. Plant hairs often produce thick secondary walls as, for instance, the cotton seed hairs or the
chamber hairs of Hamulus. The walls or trichomes are sometimes impregnated with silica or calcium
carbonate. Their contents are varied in relation to function. Cystoliths and other crystals may develop
in hairs.
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Review
Trichomes are defined as unicellular or multicellular appendages, which are an extension of the
above-ground epidermal cells in plants [1]. These appendages play a key role in the development of
plantsand occur in a wide variety of species [2]. Trichomes are a protective barrier against natural
hazards, such as herbivores, ultraviolet (UV) irradiation, pathogen attacks, excessive transpiration,
seed spread, and seed protection. The unicellular non-glandular trichomes of Arabidopsis serve as an
excellent model to study the molecular mechanism of trichome development in plants [3,4,5,6,7].
Despite the small size of trichomes, they have a marked effect on plants and human health. Cotton
seed trichomes are important raw materials in the textile industry [8,9,10], and tea trichomes are
critical for tea breeding and tea quality as they are rich in nutrients [11]. The light-leaf and light-shell
phenotypes of crop rice have missing trichomes, which are conducive to crop harvesting and
subsequent processing; however, the leaf trichomes
in Populus and Platanus are harmful to humans, easily causing respiratory tract infections, lung
infections, fever, influenza, and in severe cases, potentially cause cancer [12].
Baillon (1864, 1958) and Müller (1866, 1973) were the first botanists to explore trichome
morphology in the genus for taxonomic purposes. However, they limited their scope to two basic
types: the stellate trichome, formed by free rays, and the lepidote trichome, with rays united in
various degrees. A few decades later, Froembling published the first comprehensive study of
trichomes of the tribe Crotoneae and other groups of Euphorbiaceae s.l., including
132 Croton spp. (Froembling, 1896) and proposed a classification recognizing five basic trichome
types: lepidote (schilghaare); stellate (sternhaare); fasciculate or rosulate (buschelhaare); fasciculate-
stipitate (morgensternförmig Haare) and dendritic (candelaberhaare).
A century later, Webster et al. (1996) revisited and revised Froembling’s work. They surveyed
trichome morphology from 120 Croton spp. and proposed a new classification that included seven
basic types of trichomes: lepidote, stellate, fasciculate, multiradiate/rosulate, dendritic, papillate and
glandular. Since its publication, Webster’s classification has served as the main reference for
trichome typology and terminology, not only in taxonomic and anatomical studies focusing on
Croton (Van Ee et al., 2011; Caruzo and Cordeiro, 2013; Vitarelli et al.,
2015, 2016; Feio, et al., 2018a; Feio et al., 2018b; Riina et al., 2021), but also in other groups of
Euphorbiaceae
12

(Cervantes et al., 2009; Bigio and Secco, 2012; Riina et al., 2014; Silva and Cordeiro, 2020) and other
plant families (Clark et al., 2010; Tan et al., 2012; Wurdack, 2017; Ganesan et al., 2021).
Structure and morphology characteristics of trichomes
Trichomes are widely distributed on the surface of different organs/tissues in different plants,
exhibiting various morphologies. Trichomes are generally divided into single-celled or multicellular,
branched or unbranched, and glandular or non-glandular based on different characteristics and
functions. Trichomes also have different shapes, such as head, star, hook and scale. Theobald and
Barthlott further divided trichomes into three categories based on the distributions of leaves: large,
small and glandular trichomes [13, 14]. Large trichomes are mainly distributed on the abaxial surface
and margins and in the vascular bundles; small trichomes in the stomatal para cellular; and glandular
trichomes are usually regularly distributed in all or part of the subepidermal tissue on the leaf surface
[2]. The morphology of trichomes is associated with the spatial distribution of plant organs. The base
of the tomato (Lycopersicon esculentum) stem is well-covered with long trichomes, whereas the
upper parts of the stem has short and sparse trichomes [15]. Moreover, the density of trichomes
varies in different organs. The density of trichomes on the leaf blade is significantly higher than that
on the back of the blade [16]. Brewer found that different types of leaves had different densities of
trichomes. In soybean, the adaxial surface of the leaf had a greater trichome density than the abaxial
surface [17].
Biological functions of plant trichomes
Plants have evolved many defense mechanisms to protect against different abiotic and biotic
stresses. The morphology and density of trichomes influence several aspects of plant physiology and
ecology by mediating the interactions between the plant and its environment [18]. Trichomes, along
with the stomata, cutin and wax on the epidermis, performed various protective functions through
synthesizing, storing and secreting many important substances [19,20,21]. The cotton petal trichomes
maintain the shape of the buds and ensure the production of seeds [22]. In addition, trichomes protect
plants from herbivores, insects and pathogens by secreting repellents, alkaloids and toxic substances
[21, 23,24,25]. Kim et al. (2011) examined the relationship between pepper trichomes and pepper
mottle virus (PepMov) resistance and showed that the resistance to PepMoV-SNU1 strain is inherited
by cross combinations among pepper cultivars CM334, Chilsungcho and ECW123R [15]. In rice, two
TRICHOME BIREFRINGENCE (TBR)-like proteins play an essential role in the resistance to leaf
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spot disease [26].
Trichomes are also important in the response to abiotic stress [22]. The presence of trichomes
increases the thickness of the epidermis, and the content of long-chain fatty acids is significantly
higher than that in other epidermal cells, which is helpful to reduce evaporation and regulate
temperature [20, 27]. In Brazil’s high-altitude rocky areas, the plants of Croton tiglium and Vriesea
effectively absorb moisture and nutrients from the atmosphere through trichomes to improve water
and fertilizer utilization [28, 29]. The high-density multi-branched Acanthophyllum squarrosum
trichomes not only have high resistance to sand burial, but also reduce mechanical damage by wind
and sand [30]. In addition, the trichomes of an aquatic plant, Salvinia molesta, play a hydrophobic
role in maintainin

normal respiration [14]. Trichomes also function in signal transmission. The cell wall of Arabidopsis
trichomes gradually thins from the top to bottom, and this change makes the base of the trichomes
extremely sensitive to external stimuli. The stimulation is transmitted to cells around the base of
trichomes through changes in Ca 2+ content and pH, thereby regulating the synthesis of defensive
substances [31, 32].

Regulatory mechanism of trichome development in plants

The development of plant trichomes is coordinated and regulated by a variety of factors, such as
the environment, hormones, regulatory genes, and non-coding RNA. Among them, regulatory genes,
including transcription factors and functional genes, play important roles in regulation of the
initialization, growth and development of trichomes.

Research on trichomes in tea plants (C. sinensis)

The molecular mechanism of trichome formation has also been investigated in other model plants,
such as C. sativus [12], L. esculentum [13], B. campestris [14] and B. napus [15]. The recent study
had revealed that another regulatory mechanism might be involved in regulating trichome
development in plants. Trichome (also referred to as ‘háo’ in tea) was generally regarded as one of
the quality standards of tea due to secrete various secondary metabolites, such as flavonoids, tea
polyphenols, and amino acids. The number of tea trichomes depends on degree of tenderness of tea
14
leaves. The distribution of trichomes is especially thick on newly budded leaves and gradually
decreases as the leaf develops.
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Material and Method

1. Collection of plant samples
2. Study of Dermal tissue
3. Characterization of trichomes
Ten commonly cultivated varieties of tomato, Solanum lycopersicum, namely, Big beef, Black
prince, Supersweet, Tidy treats, Pink tiger, Sungold, Pink berkeley, Purple bumble bee, Indigo cherry
drops and Black krim were used for this experiment. Due to the extensive use of scanning electron
microscopy required, we had to limit the varieties to from a long list of all commercial varieties.

Characterization of trichomes
To assess the morphology of trichomes, images were taken at varied magnifications, mostly from
100×–2000×, to get maximum resolution. Moreover, images of certain trichomes were captured at
different angles to get more specific details. The trichomes were not only classified into glandular and
non-glandular, but more finely into sub-types based on the classification given by Watts and Kariyat
2021a. The glandular trichomes were mainly classified according to the shape and structure of their
glandular head.

After imaging, the trichome density of these varieties was assessed, by capturing the images at 100×
magnification (Fig. 1). The magnification was kept consistent among all the leaf samples. By using
“Nanoeye” software in DSEM, we measured the surface area at 100× (translates to 1.89 mm2). The
number of trichomes were counted separately according to their classification, and the trichome
density per mm2 was calculated using the equation (Watts and Kariyat 2021a):
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Result
whether intraspecific variation through domestication affects defense traits, we examined in detail,
the trichomes of 10 commonly cultivated tomato varieties. We found that there is a tremendous
variation in trichome morphology, type, density as well as dimensions among these varieties, possibly
for the first time. We also found that the density of glandular trichomes is higher than non-glandular
trichomes. Also, among glandular trichomes, subtypes were found based on the shape and structure of
their glandular head. However, the small glandular hair with large quadricellular head is most
common which was found in all the varieties. Our detailed examination adds another line of evidence
for designing trichome studies to consider this variation in trichome traits (Watts and Kariyat 2021a).
Our results are also consistent with the results of Van Dam et al. (1999), who found significant
variation in the phenotype of trichomes among the populations of Datura wrightii, another species in
Solanaceae, commonly used in ecological studies (Nunez-Farfan and Dirzo 1994; Valverde et al.
2001). And we recently (Watts and Kariyat 2021a) documented and described a well-defined
classification of trichome types from 14 wild and cultivated species of Solanum genus. However,
since, of all the cultivated Solanum species, tomato is arguably, the most important species, and this
study revealed additional results including the presence of rare glandular types trichome type
(Acuminate glandular hair with bicellular stalk and small glandular tip) in a few varieties.
Out of all the 9 types of trichomes found in tomato, 6 were glandular trichomes. The glandular
trichomes are reported to secrete many volatile compounds which are known to trigger insecticidal
effects in the plants. For example, the tomato mutant having very less glandular trichomes with
quadricellular head was found to accumulate lower amount of monoterpenes, sesquiterpenes, and
flavonoids. Consequently, this mutant was highly susceptible to the herbivores in natural conditions,
suggesting the anti-herbivore nature of these trichome secreted compounds (Kang Liu et al. 2010).
Many scientists have done the gene expression profiling of trichomes (Cui et al. 2011) and
transcriptomics (Balcke et al. 2017). Our study showed the enormous variation in the density of this
trichome along with diameter of glandular quadricellular head and can act as a foundation to decipher
the function of its size and density on the plant defense as this trichome secretes many important
compounds, possibly involved in plant defense against herbivores.
17
Contrary to the glandular trichomes, the non-glandular trichomes are documented to affect feeding
and oviposition
of herbivores because of their structure. For example, Tian et al. 2012 found that in tomato, the
mutants with high density of non-glandular trichomes negatively affect the feeding and growth of
Colorado potato beetle, Leptinotarsa decemlineata. Apart from this, the non-glandular trichomes are
found to cause post-ingestive damage by rupturing the peritrophic gut membrane of caterpillars
(Kariyat et al. 2017). Similarly, in rice the silicified non-glandular trichomes were found to damage
the gut membranes of rice chewing herbivores and thus acting as an important defense component
(Andama et al. 2020). In this study, we examined thoroughly the length variations of important non-
glandular trichomes of tomato and found that they significantly differ from each other-suggesting
potential functional consequences for both settling and feeding disruptions (Kariyat et al. 2018), and
post ingestive effects on the gut lining (Kariyat et al. 2017). Clearly, a detailed examination of diets
supplemented with trichomes from these varieties, and their effects on herbivore fitness is warranted.
Trichomes, in general, have been very well studied for their role in plant defense against herbivores
(Kariyat et
al. 2017, 2019; Watts and Kariyat 2021b). They are also highly inducible through herbivory (Kariyat
et al. 2013; Traw and Dawson 2002)), water stress (Bosu and Wagner 2014; Gonzáles et al. 2008),
and also through mechanical damage (Gonzáles et al. 2008), have differential effects on different
herbivores (Karley et al. 2016; Tozin et al. 2017), vary across leaf surfaces (Karabourniotis et al.
1999; Watts and Kariyat 2021b), and have multiple modes of action (Kaur and Kariyat 2020). Studies
like this that shows significant variation for trichomes traits (within just 10 varieties) through
morphological assays and diversity estimation can further assist in the breeding programs for
developing host resistant varieties of tomato, targeting specific pests and their life stages.

We found significant variation for trichome density in our study, not only in major trichome
types, but also in the subtypes of trichomes. The importance of trichome density variation has been
documented in many studies for its impact not only in biotic stresses (Handley et al. 2005;
Valverde et al. 2001) but also in abiotic stresses, such as protection against UV radiations
(Karabourniotis et al. 1999), protection against high insolation (Karabourniotis et al. 1999),
drought tolerance (Zhang et al. 2020), lowering transpiration water losses (Pérez-Estrada et al.
2000) and resistance to low temperature (Zhang et al. 2020). All these studies show that the
18
trichome density is significantly correlated with plant resistance to biotic and abiotic stresses.
Zhang et al. (2020) studied different mutants and cultivars of tomato with varied trichome densities
for its relationship to the stress resistance of the plants with treatments of disease, insects, drought
and cold. However, the role of density variation of different subtypes of trichomes of tomato in
stress resistance needs further investigation, especially from an abiotic stress perspective, for
drought management and climate smart crops—to build upon from our work.
The Shannon diversity index and equitability of the trichomes in different varieties was calculated.
The equitability of eight out of ten varieties came out as more than 0.7, suggesting that these varieties
are very diverse regarding their trichomes. Interestingly, the two varieties having maximum total
trichome numbers have the lowest equitability suggesting that the maximum trichome number does
not always means maximum diversity. However, the role of this

trichome density vs trichome diversity with regard to plant defense mechanisms warrants more
experiments.
Taken together, tomato is a very good model for studying the role of trichomes in plant defense as it
contains a highly diverse and varied density and dimensions of different trichomes. Future research
should explore the importance of density and dimension variation of important subtype of trichomes
in tomato. Since tomato is mainly a glandular trichome enriched species, the diversity of volatiles
from different cultivars and varieties should be explored in detail. Conclusion:
The overall goal of this study was to examine in detail, the morphological characteristics of the
trichomes of tomato, since it is one of the most important domesticated species in Solanaceae. We
found enormous variation in not only the number of total trichomes as well as their subtypes, but also
in its dimensions, showing intraspecific
variations in trichome traits in commonly cultivated varieties of tomato. This study, we believe will
lead to future work in understanding the functional consequences of this intraspecific trichome
variations in plant defense against biotic as well as abiotic stresses, by taking into consideration each
trichome subtype.
Trichomes have a marked effect on plants and human health, its growth and development are
effected by environment factors, hormones, transcription factor, non-coding RNA, etc. Although
several previous studies had summarized the factors that affected the formation and development
19
of plant trichomes, their reviews lacked a comprehensive summary and only focused on
transcription factors, hormones and environment in model plants. The aim of this review is not only
to summarize the factors, such as environment factors, hormones, transcription factor, non-coding
RNA, and functional genes, regulate the formation and development of plant trichomes in model
plants, but also investigate the regulation mechanism of trichome development in non-model
plants.
20

Conclusions

Trichomes have a marked effect on plants and human health, its growth and development are
effected by environment factors, hormones, transcription factor, non-coding RNA, etc. Although
several previous studies had summarized the factors that affected the formation and development of
plant trichomes, their reviews lacked a comprehensive summary and only focused on transcription
factors, hormones and environment in model plants. The aim of this review is not only to summarize
the factors, such as environment factors, hormones, transcription factor, non-coding RNA, and
functional genes, regulate the formation and development of plant trichomes in model plants, but also
investigate the regulation mechanism of trichome development in non-model plants, such as tea. We
are expected this review provided a reference for basic and applied research on trichomes in plants in
the near future.
21

References
1. Johnson HB. Plant pubescence: An ecological perspective. Bot Rev. 1975;41(3):233–58.
2. Xiao K, Mao XH, Lin YL, Xu HB, Zhu YS, Cai QH, et al. Trichome, a functional diversity

phenotype in plant. Mol Biol. 2016;2017(1):1–6.

3. Lloyd A, Schena M, Walbot V, Davis R. Epidermal cell fate determination in Arabidopsis:

patterns defined by a steroid-inducible regulator. Science. 1994;266(5184):436–9.

4. Chien JC, Sussex IM. Differential regulation of trichome formation on the adaxial and abaxial

leaf surfaces by gibberellins and photoperiod in Arabidopsis thaliana (L.) Heynh. Plant
Physiol. 1996;111(4):1321–8.
5. Larkin JC, Young N, Prigge M, Marks MD. The control of trichome spacing and number

in Arabidopsis. Development. 1996;122(3):997–1005.

6. Telfer A, Bollman KM, Poethig RS. Phase change and the regulation of trichome distribution

in Arabidopsis thaliana. Development. 1997;124(3):645–54.

7. Perazza D, Vachon G, Herzog M. Gibberellins promote trichome formation by up-regulating

GLABROUS1 in Arabidopsis. Plant Physiol. 1998;117(2):375–83.

8. Wang G, Feng HJ, Sun JL, Du XM. Induction of cotton ovule culture fibre branching by co-
expression of cotton

BTL, cotton SIM, and Arabidopsis STI genes. J Exp Bot. 2013;64(14):4157–68.
9. Guan XY, Pang MX, Nah G, Shi X, Ye W, Stelly DM, et al. miR828 and miR858 regulate
homoeologous MYB2 gene functions in Arabidopsis trichome and cotton fibre development.
Nat Commun. 2014. https://doi.org/10.1038/ncomms4050.
10. Li WQ, Wu JG, Weng SL, Zhang D, Zhang Y, Shi C. Characterization and fine mapping of the

glabrous leaf and hull mutants (gl1) in rice (Oryza sativa L.). Plant Cell Rep. 2010;29(6):617–
27.
11. Li JL, Zeng LT, Liao YY, Tang J, Yang Z. Evaluation of the contribution of trichomes to

metabolite compositions of tea (Camellia sinensis) leaves and their products. LWT Food Sci
Technol. 2020.
12. Tang XL, You T, Wu W, Sun L, Li Z. Allergenic property and two kinds of common street trees in

Nanjing and the control. J Jishou Univ (Natural Science Edition). 2014;35(3):69–73.
22
13. Theobald WL, Krahulik JL, Rollins RC. Trichome description and classification. Anatomy of

the dicotyledons I. Oxford: Clarendon Press; 1980. p. 40–53.

14. Barthlott W, Wiersch S, Colic ZC, Koch K. Classification of trichome types within species

of the water fern Salvinia, and ontogeny of the egg-beater trichomes. Botany.
2009;87(9):830–6.
15. Kim HJ, Han JH, Kim S, Lee HR, Shin J, Kim J, et al. Trichome density of main stem is tightly

linked to PepMoV resistance in chili pepper (Capsicum annuum L.). Theor. Appl. Genet.
2011;122(6):1051–8.
16. Ascensão L, Marques N, Pais MS. Glandular trichomes on vegetative and reproductive organs of

Leonotis leonurus (Lamiaceæ). Ann Bot. 1995;75(6):619–26.

17. Brewer CA, Smith WK. Influence of simulated dewfall on photosynthesis and yield in soybean

isolines (Glycine max [L.] merr. cv Williams) with different trichome densities. Int J Plant Sci.
1994;155(4):460–6.
18. Wagner GJ, Wang E, Shepherd RW. New approaches for studying and exploiting an old

protuberance, the plant trichome. Ann Bot. 2004;93(1):3–11.

19. McDowell ET, Kapteyn J, Schmidt A, Li C, Kang J, Descour A, et al. Comparative functional

genomic analysis of Solanum glandular trichome types. Plant Physiol. 2010;155(1):524–39.

20. Hegebarth D, Buschhaus C, Wu M, Bird DA, Jetter R. The composition of surface

wax on trichomes of Arabidopsis thaliana differs from wax on other epidermal cells.
The Plant J. 2016;88(5):762–74.
21. Rakha M, Bouba N, Ramasamy S, Regnard J, Hanson P. Evaluation of wild tomato accessions

(Solanum spp.) for resistance to two-spotted spider mite (Tetranychus urticae Koch) based on
trichome type and acylsugar content. Genet. Resour. Crop Ev. 2017;64(5):1011–22.
22. Zhao Q, Chen XY. Development: a new function of plant trichomes. Nat Plants. 2016;2(7):16096.
23. Kang JH, Liu G, Shi F, Jones AD, Beaudry RM, Howe GA. The tomato odorless-2 mutant is

defective in trichome- based production of diverse specialized metabolites and broad-spectrum

resistance to insect herbivores. Plant Physiol. 2010;154(1):262–72.
24. Kang J, Shi F, Jones AD, Marks MD, Howe GA. Distortion of trichome morphology by the hairless
mutation of

tomato affects leaf surface chemistry. J Exp Bot. 2009;61(4):1053–64.

23
25. Maes L, Van Nieuwerburgh F, Zhang Y, Reed DW, Pollier J, Casteele SV, et al. Dissection of the

phytohormonal regulation of trichome formation and biosynthesis of the antimalarial compound

artemisinin in Artemisia annua plants. New Phytol. 2010;189(1):176–89.
26. Gao Y, He C, Zhang D, Liu X, Xu Z, Tian Y, et al. Two trichome birefringence-like

proteins mediate xylan acetylation, which is essential for leaf blight resistance in rice. Plant
Physiol. 2017;173(1):470–81.
27. Busta L, Hegebarth D, Kroc E, Jetter R. Changes in cuticular wax coverage and composition on
developing Arabidopsis leaves are influenced by wax biosynthesis gene expression levels and
trichome density. Planta. 2017;245(2):297–311.
28. Vitarelli NC, Riina R, Cassino MF, Meira RM. Trichome-like emergences in croton of

Brazilian highland rock outcrops: evidences for atmospheric water uptake. Perspect Plant Ecol.
2016;1:23–35.
29. Vanhoutte B, Schenkels L, Ceusters J, De Proft MP. Water and nutrient uptake in vriesea

cultivars: trichomes vs. roots. Environ Exp Bot. 2017;1:21–30.

30. Chen G, Zhao J, Zhao X, Zhao P, Duan R, Nevo E, et al. A psammophyte Agriophyllum

squarrosum (L.) Moq.: a potential food crop. Genet. Resour. Crop Ev. 2014;61(3):669–76.
31. Liu H, Zhou LH, Jiao J, Liu S, Zhang ZM, Lu TJ, et al. Gradient mechanical

properties facilitate Arabidopsis trichome as mechanosensor. Appl Mater

Interfaces. 2016;8(15):9755–61.
32. Zhou LH, Liu SB, Wang P, Lu TJ, Xu F, Genin GM, et al. The Arabidopsis trichome is an active
mechanosensory switch. Plant Cell Environ. 2017;40(5):611–21.
33. Oppenheimer DG, Herman PL, Sivakumaran S, Esch JJ, Marks MD. A MYB gene required

for leaf trichome differentiation in Arabidopsis is expressed in stipules. Cell.

1991;67(3):483–93.
34. Dai X, Wang G, Yang DS, Tang Y, Broun P, Marks MD, et al. TrichOME: a comparative omics

database for plant trichomes. Plant Physiol. 2009;152(1):44–54.

35. Alahakoon UI, Taheri A, Nayidu NK, Epp DJ, Yu M, Parkin IA, et al. Hairy canola (Brasssica

napus) re-visited: down-regulating TTG1 in an AtGL3-enhanced hairy leaf background improves

growth, leaf trichome coverage, and metabolite gene expression diversity. BMC Plant Biol.
2016;16(1):e12.
24
36. Ioannidi E, Rigas S, Tsitsekian D, Daras G, Alatzas A, Makris A, et al. Trichome patterning

control involves TTG1 interaction with SPL transcription factors. Plant Mol Biol. 2016;92(6):675–
87.
37. Li Y, Shan X, Gao R, Yang S, Wang S, Gao X, et al. Two iiif clade-bHLHs from Freesia

hybrida play divergent roles in flavonoid biosynthesis and trichome formation when ectopically
expressed in Arabidopsis. Sci Rep. 2016;6(1):e30514.
38. Gao C, Li D, Jin C, Duan S, Qi S, Liu K, et al. Genome-wide identification of GLABRA3

downstream genes for anthocyanin biosynthesis and trichome formation in Arabidopsis.

Biochem Bioph Res Co. 2017;485(2):360–5.
39. Johnson CS, Kolevski B, Smyth DR. TRANSPARENT TESTA GLABRA2, a trichome and seed
coat development
gene of Arabidopsis, encodes a WRKY transcription factor. Plant Cell. 2002;14(6):1359–75.
40. Dai X, Zhou L, Zhang W, Cai L, Guo H, Tian H, et al. A single amino acid substitution in

the R3 domain of GLABRA1 leads to inhibition of trichome formation in Arabidopsis

without affecting its interaction with GLABRA3. Plant Cell Environ. 2016;39(4):897–907.
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JAVA WHITE PLANT

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PURPLE HEART PLANT

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