Xerophytes, roughly translating to “dry plants” ​(Delf 1915), ​have undergone special

modifications and adaptations to be able to survive in an environment with little availability to

water or moisture. While most xerophytes are commonly found in the arid conditions of the

desert, few xerophytes may also be found in tropical habitats where water supply is limited or

too intermittent for mesophytic plants ​(“Desert Plant Survival” 1996)​. On top of this, plants

living in arctic conditions may also require xerophytic adaptations as water is unavailable for

uptake when the ground is frozen ​(“Desert Plant Survival” 1996)​. In order to survive, xerophytes

have adopted a variety of techniques that are critical to their survival and growth under various

environmental pressures-aiming to counter the water deficiency characterized in a xerophytic

habitat. The adaptations xerophytes undergo can be classified into 3 categories: adaptations to

water loss, adaptations to store water, and adaptations for water uptake. These specialized

modifications allow xerophytes to tolerate extremely dry conditions, low humidity, high

wind-flow, and water droughts ​(“Desert Plant Survival” 1996)​.

The modifications in the leaves of xerophytes perform crucial water conservation

strategies ​(Beaumont 2014)​. The reduced surface area of xerophytic leaves are usually

accompanied by changes in the leaf’s internal structure that have adapted to provide protection

against leaf desiccation and towards water retention ​(Fatima et al 2018)​. Since the preservation

of water is of utmost importance for plants growing in arid environments, xerophytes tend to

have small, waxy leaves, which help the plant to retain water ​(Fatima et al 2018)​. The first line

of defense in response to water scarcity is the thickening of the epidermis and cuticle to

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minimize water loss from the plant body ​(Liesenfeld et al 2019)​. Since most water loss in plants

occurs through transpiration from leaves, desert plants have modified their leaves into needles,

and in some cases, have even abandoned their leaf structures entirely in order to reduce the

surface area of the leaf ​(Liesenfeld et al 2019)​. The sharp spines that you see on the cactus and

other plants further aid in the reduction of transpiration by providing shade for the plant from the

sun ​(Beaumont 2014)​.​ ​In addition, the overall small stature of xerophytic plants demonstrate how

stunted growth can be extremely beneficial for water conservation in times of drought, as plants

use their energy towards means of survival, rather than for normal vegetative growth ​(Mansoor

et al 2015). ​The rate of transpiration through leaves are also controlled by stomatal pore size,

density, and orientation ​(Liesenfeld et al 2019)​. Desert plants generally have smaller and lesser

stomata, located on the abaxial surface of the leaf. The location of the stomata removes them

from direct sunlight, keeping the gas exchange surface cool ​(Liesenfeld et al 2019)​. Moreover,

smaller stomata may regulate the rate of transpiration more effectively than larger stoma as less

water is required for turgor maintenance ​(Liesenfeld et al 2019)​, and thus, protecting plants from

desiccation when water availability is limited. In addition to the thicker epidermal layer of leaves

and stomatal pores, epidermal appendages are amongst the most important characteristics that

protect plants from ​environmental​ stresses. The dense long hairs (trichomes) of desert flora help

control the temperature of the leaf and minimize water loss through their leaf’s surfaces as they

reflect solar radiation, resist wind speed, and develop a humid layer near the leaf's surface

(Prasad 1996)​. Moreover, trichomes are also impermeable to water, and therefore, protect leaves

from undue water loss ​(Prasad 1996)​. Another feature related to transpiration is the adaptation of

bulliform cells. These cells aid in the conservation of water through their ability to roll their

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leaves​,​ as well as through their ability to store greater quantities of water ​(Hernandez et al 2016)​.

Under stress, such as during long periods of drought, the bulliform cells enable the leaves to roll

unto themselves and then unwrap when rehydrated ​(Hernandez et al 2016)​. While rolled, sun

exposure to the surface area and shoot are limited, reducing the rate of transpiration significantly

(Hernandez et al 2016)​.

Storage parenchyma and cortical cell areas in xerophytes serve an essential role in storing

water and nutrients ​(“Xerophytes” 2016)​. An increase in succulence resulting from the storage

parenchyma enables the plant to conserve water inside plant tissues, whereas the cortical cell

area produces exceptionally high root and leaf sheaths, as well as larger vacuoles ​(Beaumont

2014)​. These features combined, work together to provide a greater capacity for the storage of

water and minerals to be used during ​drought​ periods ​(Beaumont 2014)​, hence, making the plant

more resistant to unfavourable conditions. Many species of xerophytes have adapted to desert

conditions through the development of extensive root systems. Some species have root systems

that draw water from deep under the soil, encouraging maximum absorption of what little water

is available ​(Bibi et al 2015)​. On the other hand, the cactus displays a shallow root system that

allows for the quick acquisition of large quantities of water in brief rainy periods ​(Bibi et al

2015)​. The thick stems of cacti are a result of their ability to store more than five tonnes of water

in the core of both stems and roots, providing them with the resources to survive in years of

drought on the water collected from a single rainfall ​(Mansoor et al 2015)​. In addition, the

Creosote bush withholds a double root system that allows them to accumulate water from both

surface and groundwater ​(Dimmitt 1997)​.

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 Due to the harsh environmental extremities of xerophytic habitats, most seeds go through

a resting period or dormancy. In order for seeds to germinate, certain environmental conditions

must be present-only then can a seed germinate and develop into a new plant ​(Lai et al 2016)​.

The length of seed dormancy varies greatly, with some dormancy periods extending from days to

years, as the embryo cells in the seed metabolize very slowly ​(Lai et al 2016).​ Desert plant seeds

must only germinate when there is enough moisture in the ground to support their growth to

maturity. The hard and impermeable seed coat allows the seeds to endure the stressful high

temperatures ​(Lai et al 2016)​. Xerophytic plants have a chemical in their seeds that inhibit

germination ​(MacMillan 1978)​. Only heavy rainfall is able to wash away this inhibitor and let

the seed germinate, a light rain is not enough to break their dormancy ​(MacMillan 1978)​. If they

germinate after a brief rain, the seedling could dry out and die after only a week or so ​(Lai et al

2016)​. As the long dry periods begin, only the seeds survive until the next period of rain arrives,

while the plants develop rapidly during the short rainy season, producing flowers and seeds

within a few weeks ​(Lai et al 2016)​. Desert perennials, such as the Ocotillo, often survive by

becoming dormant during dry periods of the year, then springing to life when water becomes

available ​(“Seed Dormancy” 2020)​. After rain falls, the Ocotillo quickly grows new leaves to

photosynthesize food ​(“Seed Dormancy” 2020)​. The flowers bloom within a few weeks, and

when seeds become ripe and fall, the Ocotillo loses its leaves again and re-enters dormancy

(“Seed Dormancy” 2020)​. Because this process may occur as many as five times a year, the

Ocotillo also has a waxy coating on stems that serves to seal in moisture during these periods of

dormancy ​(“Seed Dormancy” 2020)​. Another example of desert perennials that utilize seed

dormancy to survive drought are the bulb members of the lily family. They are able to store

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enough nourishment to survive for long periods in rocky or alluvial soils ​(Lai et al 2016)​. The

tops of bulbs dry out completely, leaving no trace of their existence above ground during

dormant periods ​(Lai et al 2016).

Xerophytes have adapted to these dry and arid habitats through behavioural and

morphological modifications, allowing them to better conduct and conserve water during times

of drought. These adaptations all work together to ensure the survival of xerophytes, despite the

environmental pressures they face.