ISSN 1019-3316, Herald of the Russian Academy of Sciences, 2017, Vol. 87, No. 1, pp. 1–11.

© Pleiades Publishing, Ltd., 2017.

Original Russian Text © N.Z. Shamsutdinov, E.Z. Shamsutdinova, N.S. Orlovsky, Z.Sh. Shamsutdinov, 2017, published in Vestnik Rossiiskoi Akademii Nauk, 2017, Vol. 87, No. 1,
pp. 3–14.

Science and Society

Halophytes: Ecological Features, Global Resources,

and Outlook for Multipurpose Use
N. Z. Shamsutdinova1, E. Z. Shamsutdinovab2, N. S. Orlovskyc3, and Z. Sh. Shamsutdinovb4, *
a
Kostyakov All-Russian Research Institute of Hydraulic Engineering and Land Reclamation, Moscow, Russia
b
Vil’yams All-Russian Fodder Research Institute, Lobnya, Russia
c
Blaustein Institutes for Desert Research, Ben-Gurion University of the Negev, Sede Boker, Israel
e-mail: 1nariman@vniigim.ru; 2darplant@mtu-net.ru; 3nicolai@bgu.ac.il; 4aridland@mtu-net.ru
Received February 20, 2016

Abstract—Halophytes (galos, salt; phyton, a plant) are ecologically, physiologically, and biochemically spe-
cialized plants, which belong to various systemic and life forms capable of finishing their life cycle in a saline
habitat (the soil solution’s electrical conductivity is 8–10 dS/m). Halophyte resources are an important
source and reserve of the sustainable development of agriculture in arid parts of the world, including Russia.
The importance of the world’s halophyte resources is shown for obtaining fodder, grass forage, and medicinal
and oil raw materials, as well as biological agents for reclaiming degraded lands, especially in arid regions,
where dire shortages of food are observable. Research results of introducing halophytes in crops as forage, oil,
and medicinal plants in the conditions of irrigated and nonirrigated farming in arid regions of Central Asia
and Russia are given. Prospects for the multipurpose use of halophytes in the sustainable development of via-
ble agriculture are substantiated.

Keywords: halophytes, resources, salt resistance, irrigation, salt waters, multipurpose use, arid zone, intro-
duction of halophytes in crops.
DOI: 10.1134/S1019331616060083

At present, one of the most important problems of
humanity is the search for sufficient quantities of water
and land resources to meet the need for food. Accord-
ing to the data of the UN Food and Agriculture Orga-
nization, it is necessary to bring 200 mln hectares of
land under cultivation in the next 30 years to feed the
growing population of the planet. Thus far, there are
only 93 mln hectares, but they require deforestation
and preparatory measures. Alternative phytore-
sources, as well as water and land resources, should be
found to expand the area of cultivated crops [1].
Among the species of natural flora, there are eco-
logically, biologically, physiologically, and biochemi-
cally specialized plant organisms capable of function-
ing and reproducing normally on saline soils and/or
* Nariman Zebrievich Shamsutdinov, Dr. Sci. (Biol.), is Head of
the Laboratory of Phytomelioration of Degraded Lands at the
Kostyakov All-Russian Research Institute of Hydraulic Engi-
neering and Land Reclamation. El’mira Zebrievna Shamsutdi-
nova, Cand. Sci. (Agric.), is Head of the Department of Arid
Forage Plants at the Vil’yams All-Russian Fodder Research
Institute. Nikolai Sergeevich Orlovsky, Dr. Sci. (Geogr.), is a
Professor at the Blaustein Institutes for Desert Research, Ben-
Gurion University of the Negev. RAS Corresponding Member
Zebri Shamsutdinovich Shamsutdinov is Head of the Selection
Center, Vil’yams All-Russian Fodder Research Institute.
when irrigated by salt water [2]. These plants are called
halophytes.
The range of salinity of the soil solution within
which a plant can grow and reproduce normally is dif-
ferent in various halophyte species. Over the past 20–
25 years, the attention of scientists has been aimed at
the study and distribution of halophytes to produce
food (grains, grass forage, feeds, oil and medicinal raw
materials) in arid parts of the world based on the use of
salt water for irrigation (sea, ground, and collector-
and-drainage waters). This problem is topical for most
countries, because freshwater resources are scarce,
only 2.53% of the global water resources. At the same
time, salt water reserves are huge, 97.47% [3]. As the
consumption of freshwater increases, its deficit will
escalate significantly in the next few years in all coun-
tries of the world. As the earth’s population is con-
stantly growing, the food situation is becoming worse
year after year. These circumstances dictate the need
to seek ways of using salt water to stimulate food pro-
duction using halophytes [1, 2].
Abroad, great attention to raising halophytes on
salinized soils of the arid zone was generated after the
publication in the 1960s of the results of very interest-
ing experiments in irrigating plants with undiluted sea-
water, which were conducted in Israel [1].

1
2 SHAMSUTDINOV et al.
The global scientific community, concerned with
the critical state of food resources, referred to the
potential possibilities of using the great salt water
resources of seas and oceans, as well as ground and
collector-and-drainage waters, to organize food pro-
duction. This issue is currently in the focus of UNE-
SCO and the European Union. The new technology
of cultivating halophytes using salt water, which has
been developed in a number of countries, may become
an important source of food in the near future [4].
Taking into account the global importance of rais-
ing halophytes, the IAEA has undertaken to finance
research within the interregional project “Sustainable
Use of Saline Groundwater and Wastelands for Plant
Growing” in Egypt, Iraq, Morocco, Pakistan, Syria,
Algeria, Jordan, Italy, and Germany. Almost all coun-
tries plan to share the results of their studies with other
regions by creating national halophyte programs. At
present the project’s member countries are able to
grow halophytic plants useful for the economy on sali-
nized nonbearing wastelands using saline waters.
Halophyte farming, as global and domestic experi-
ence shows, solves a triune problem: the economy of
freshwater and finances; the reclamation of the envi-
ronment, including soil fertility; and the production of
agricultural products (grains, grass forage, feeds, and
oil and medicinal raw materials) [2]. Halophyte crop
production can be organized primarily on sandy soils
in desert regions and under irrigation by saline (sea,
ground, and drainage) water. By the assessment given
in [5], about 15% of uncultivated lands that belong to
littoral and continental deserts could be suitable for
farming if irrigated by seawater, which is equivalent to
130 mln hectares of new plantings.
Solar energy and saltwater resources are most
important on vast arid territories and are still unused
for crop production in Russia and in the world in gen-
eral. In addition, there are large spaces inside the con-
tinents where brackish water reserves occur, which can
be used for highly productive crop growth [5]. The
waters of seas and oceans are the second major natural
resource for halophyte growth [1, 5].
In the opinion of authoritative ecologists [2], crop
production using freshwater for irrigation is a conse-
quence of the selection and domestication of plants
that come from ancestors that grew on freshwater.
These scientists also assert that no special reason exists
to stick to this trend. Nothing hinders the selection
and domestication of plant genotypes from natural
(wild-growing) flora tolerant to high salinity. It is pos-
sible to start seeking natural salt-tolerant halophytes
across the world and cultivating them [1, 5].
Thus, the centuries-long experience of crop pro-
duction in arid regions, oriented at using freshwater for
irrigation, represents the result of selecting and
domesticating mesoglycophyte plants, which prefer
habitats with a medium humidity. Contemporary crop
production may be characterized as mesoglycophytic.
HERALD OF THE RUSSIAN ACADEMY OF SCIENCES
Now is the time to turn to halophyte farming, develop
its concept and technology, and expand the use of
halophyte species and varieties [2]. In this context, the
most important task of modern science and agricul-
tural practice is to mobilize the genetic resources of
halophytes; create their collection; assess them eco-
logically; select valuable feed, food, medicinal, and oil
species; and domesticate them.
Among the most prominent centers where halo-
phytes are studied and domesticated for sustainable
development of profitable agriculture in arid regions
of the world are Arizona State University (United
States), Sonora State Center for the organization of
agriculture and water resources (Mexico), Ben-
Gurion University of the Negev (Israel), the Interna-
tional Center for Biosaline Agriculture (UAE), the
Vil’yams All-Russian Fodder Research Institute, and
the Kostyakov All-Russian Research Institute of
Hydraulic Engineering and Land Reclamation (Rus-
sia).
Halophyte crop production represents a major
trend in the development and use of territories hardly
suitable for agriculture due to the shortage or full
absence of freshwater. Halophytes are an alternative
source of feeds, grains, grass forage, and medicinal
and oil raw materials on lands irrigated by saline
waters, as well as a means of reconstructing the plant
cover and improving the biological productivity of
degraded pasture lands on arid territories. In order to
use this source, the Vil’yams Institute established the
Department of Arid Forage Plants and the Cis-Cas-
pian base station and the Kostyakov Institute set up
the Laboratory of Phytomelioration of Degraded
Lands.
Characteristic features of halophyte ecology. The
ability of halophytes to function normally and form a
high feed mass and medicinal raw materials in saline
conditions is due to their specific ecological, physio-
logical–biochemical, and anatomical–morphological
characteristics. All halophytes are adapted to limiting
factors of the environment, which determine the
growth and geography of distribution on arid territo-
ries, primarily to moisture shortages due to osmotic
and toxic effects of salts on plants and the physiologi-
cal dryness of soil, preconditioned by a high ion con-
tent in the soil solution [6].
Let us enumerate the major physiological–bio-
chemical and anatomical–morphological features of
halophytes that ensure their normal existence in saline
environments.
First, there is secretion of salts into the environ-
ment and the maintenance of a balanced salt content
in the cell cytoplasm. Salt-releasing halophytes are
usually nonsucculent plants with salt-releasing glands
located on photosynthesizing organs [7]. With the
help of the salt-releasing mechanism, the plants
remove surplus salts from tissues, thus regulating their
mineral composition. Limonium spp., Tamarix spp.,
Vol. 87 No. 1 2017
 HALOPHYTES: ECOLOGICAL FEATURES
and Frankenia spp. may be attributed to this group.
Other halophytes release salts with the help of salt-
accumulating vacuoles, in which the salt concentra-
tion is higher than in mesophyll cells. Another means
of salt content adjustment is the abscission of salt-
accumulating organs (Halocnemum Bieb., Halostachys
C.A. Mey, etc.) [8].
Second, some halophytes that belong to succulents
can accumulate high-concentration salts in the cellu-
lar fluid. Succulence develops at a large inflow of
chlorides, which leads to the swelling of protein and,
consequently, to ion hydration of the protoplasm.
During water absorption, this results in cell hypertro-
phy. Species of the genera Salsola L., Salicornia L.,
etc., belong to this group.
Third, a high osmotic pressure in the cells of halo-
phytes is formed due to an increase in the content of
ions and low-molecular organic compounds (pro-
lines, betaines) in cells. As a rule, water famine stimu-
lated certain biological processes in halophytes that
are ecologically significant in saline conditions. Pro-
line accumulation is among the characteristic reac-
tions [9]. It affects osmoregulation, acts as a protector
against dehydration, and serves as a source of energy
and nitrogen for metabolic processes. A high osmotic
pressure in root and shoot tissues should be considered
as an important physiological feature that helps
improve the efficiency of water absorption. The value
of osmotic pressure can serve as an accurate indicator
of water balance and, consequently, the general water
regime of a plant [6, 10]. A high osmotic pressure is
characteristic of plants that grow on salinized territo-
ries, e.g., Atriplex halimus R. Br. (35–70 atm), Limo-
niastrum guyonianum Durien (49.6–53.4 atm), and
Haloxylon aphyllum (Minkw.) Iljin (40–60 atm).
Fourth, as is known, three types of photosynthesis
are distinguished among the plant diversity of natural
and cultural flora: С3, С4, and CAM (Crassulacean
acid metabolism). The overwhelming majority of
halophytes belong to plants with С4 photosynthesis [5,
11, 12]. Such halophytes differ in the anatomical and
physiological features. The leaf mesostructure con-
sists of chlorophyll-bearing tissues of large thick-wall
cells around vascular bundles, which are surrounded
by one or several layers of loosely adjacent mesophyll
cells. Note that, in С4 plants, net photosynthesis (pho-
tosynthetic activity minus respiratory activity, mea-
sured by net absorption of carbon dioxide) usually
increases at high temperature and light intensity val-
ues. The efficacy of water utilization in С4 plants is
usually higher than in С3 plants; i.e., С4 plants require
fewer units of water to fix a СО2 unit and create a unit
of dry matter [12].
A common characteristic feature of the enzymatic
status of С4 halophytes of the goosefoot family is the
increased activity of aspartate aminotransferase
(АAТ) compared to the corresponding gramineous
groups with Kranz syndrome (also “coronary syn-
HERALD OF THE RUSSIAN ACADEMY OF SCIENCES
3
drome,” from the German Kranz, crown; in such
plants, the mesophyll cells and bundle sheathes form
two concentric layers). Within the NAD-ME and
NADP-ME types, species groups were identified that
differed significantly by the activity of enzymes of the
cycle of dicarboxylic acids [12].
According to V.I. P’yankov and A.T. Mokronosov
[13], goosefoot halophytes with С4 photosynthesis
represent plants tolerant to stress. It turns out that the
ecological role of halophytes in arid ecosystems
increases as the edaphic regime deteriorates. At the
same time, the group of goosefeet halophytes is not
homogeneous in its ecological properties. It was
shown that out of 33 С4-specific goosefoots, which
included NADM-ME and NAD-ME forms, in differ-
ent parts of the Karakum Desert, NADM-ME species
with the salsoloid and kochioid types of the Kranz
anatomy were mainly found in the most favorable con-
ditions (sands and clay sands). Areas with extreme
edaphic conditions (salinized takyrs, gypsum deserts,
puff and wet solonchaks) are populated exclusively by
NAD-ME species, among which all structural types of
the coronary syndrome of goosefoot halophytes are
found, but plants with salsoloid and suaedioid anato-
mies prevail. As ecological conditions deteriorated,
regular changes were observed in the occurrence of
biochemical types of halophytes in the direction
NADM-ME → NADP and NAD-ME → NAD-ME.
This type of photosynthesis in halophytes ensures the
normal course of the synthesis of organic substances
always with a positive balance in the conditions of the
permanent dominance of extreme factors, caused by
high temperatures, climate aridity, and soil salinity. It
follows from the above that the general evolutionary
line of halophyte adaptation by the aridity and salinity
gradients is expressed in the change in the ratio of the
photosynthesis type in ecosystems in the direction
С3 → С4 → САМ [12, 13].
Thus, the history of the origin and evolution of
halophytes (especially the goosefoot family) is closely
related to climate aridization processes and the
appearance of dry and salinized territories. Plants with
departures of metabolism and the structure of the
photosynthetic apparatus from the norm concentrate
in free ecological niches. Properties useful for plants
(the high activity of PEP carboxylase, chondriome
development, RuBP carboxylase movement in the
sheath cell, and its protection against oxygen) were
fixed during natural selection. Their integration has
led to the formation of the С4 syndrome, which is a
preadaptation and basis for the development of a wide
range of ecological niches, the formation of the spe-
cific and ecotypic diversity of halophytes in the condi-
tions of a destabilized ecological environment [12].
Fifth, anatomical–morphological features of halo-
phytes are oriented at reducing moisture consumption
by leaves and shoots. As a rule, the leaves and shoots
of halophytes are covered with a protective wax bloom
Vol. 87 No. 1 2017
4 SHAMSUTDINOV et al.

Table 1. Distribution of halophyte species by geographical E.P. Korovin [16], halophytes originated and devel-
regions oped in areas with significant concentrations of salts in
Number soil. At first, these were littorals; later, alluvial plains
Geographical distribution % and secondary salt accumulations in irrigated regions.
of species
Such ecological conditions characterize arid territo-
Asia 238 15.28 ries of Asia, Australia, America, and Africa; therefore,
Australia and Asia 8 0.51 the largest number of halophyte species concentrate
Africa 190 12.20
there.
South America 148 9.50
Most halophytes are found in the families of Che-
nopodiaceae Vent., Poaceae Barnhart, Asteraceae
Australia and Africa 5 0.32 Dumort., Plumbaginaceae Juss., Aizoaceae Rudolphi,
Australia 219 14.06 Cyperaceae Juss., Papilionaceae Giseke, Tamaricaceae
Cosmopolite 39 2.50 Link., Arecаceae Sch. Bip., and Zygophyllaceae R. Br.
North America 223 14.31
In the flora of the world, the families of Poaceae Barn-
hart (137 species), Asteraceae Dumort. (69), Plumbag-
Pantropical 36 2.31 inaceae Juss. (57), and Aizoaceae Rudolphi (53) are
Central America 60 3.85 not only exclusively halophytic but also comprise the
Sahara—Arabian and Sudanese 43 core of halophytic plant communities in various
2.76 regions of the globe.
regions
New World’s tropics 14 0.90 Analysis of halophyte flora at the generic level
shows that it includes 550 genera with the number of
Europe and North America 13 0.83
species in each of them ranging from 1 to 111. The larg-
Europe 71 4.56 est halophyte genera are given in Table 2. Among poly-
Mediterranean 70 4.49 morphic genera, we should distinguish Atriplex (111),
Old World’s tropics 67 4.30 Limonium (51), Tamarix (37), Suaeda (36), Halosarcia
(23), Sporobolus (21), Maireana (17), Frankenia (15),
America 29 1.86
and Salicornia (15); 320 genera are monotypic and
Eurasia 15 0.96 comprise 20.54% of the total number of halophytic
Australia 31 1.99 species of the world’s flora.
Africa and America 4 0.26 The species potential of halophytes of the world’s
Oceania 13 0.83 flora is represented by various life forms (Table 3):
annuals, aquatic plants, short-growing and dwarf
Australia and America 2 0.13
shrubs, thornbushes, geophyte shrubs, hemicrypto-
Africa and Asia 5 0.32 phytes, perennial grasses, sea succulents, trees, trailing
Boreal 2 0.13 and creeping plants, and parasites. Perennial grasses
Northern Hemisphere 1 0.06 prevail in the halophyte flora (25.9%); then come trees
(18.6%), shrubs (14.9%), and subshrubs with dwarf
Loss 12 0.77
subshrubs (12%).
Total 1558 100
All species of halophytes are subdivided into several
ecological groups in relation to substrate salinity and
soil texture. According to [1], hyperhalophytes com-
and have gray trichomes. Another adaptive feature prise 45.25%; xerohalophytes, 29.54%; psammohalo-
conditions a decrease in water loss by reducing the leaf phytes, 16.29%; and other groups (chasmophytes,
surface or by fully shedding leaves or part of shoots in weeds, phreatophytes), 6.35%.
the dry season of the year [14].
Positive relations exist between the type of photo-
Phytoresources of halophytes of the world’s flora. A synthesis (С4) and salt resistance, which is important
sufficiently full summary of the world’s genetic when assessing the ecological and productive capabil-
resources of halophytes is in the book [1], published in ities of plants. A series of tests in Russia and abroad
1989 in Tucson (United States). These data, as well as helped formulate a general concept of the С4 syn-
those given in [15], were processed with multidimen- drome. Evolutionarily younger taxa have it; their rep-
sional statistical methods (factor analysis, discrimi- resentatives being notable for a high temperature opti-
nant analysis, etc.), making it possible to make a con- mum of photosynthesis and a high plateau of light sat-
clusion on the distribution of halophytes by conti- uration. Over 1500 species from 18 families with this
nents; geographical regions; countries; the frequency photosynthesis type have been identified to date.
of occurrence of species, genera, and life forms; and Despite the small share in the entire diversity of higher
photosynthesis types. plants, the С4 species are of natural and economic
Analyzing the distribution of halophytes by geo- importance, especially in arid ecosystems. According
graphical regions (Table 1), note that, according to to P’yankov et al. [12, 13], with an insignificant spe-

HERALD OF THE RUSSIAN ACADEMY OF SCIENCES Vol. 87 No. 1 2017

 HALOPHYTES: ECOLOGICAL FEATURES 5

Table 2. Generic diversity of the world’s halophytic flora

Share of the total Share of the total
Number Number
Genus number Genus number
of species of species
of halophytes, % of halophytes, %
Atriplex L. 111 7.12 Prosopis Latr. 13 0.83
Liminium Mill. 51 3.27 Puccinellia Parl. 13 0.83
Tamarix L. 37 2.37 Spartina Schreb. 13 0.83
Suaeda Forssk.ex Scop. 36 2.31 Arthrocnemum Moq. 12 0.77
Halosarcia Paul G. Wilson 23 1.48 Avicennia L. 12 0.77
Sporobolus R. Br. 21 1.35 Lycium L. 12 0.77
Maireana Moq. 17 1.09 Pandanus Hemsl. 12 0.77
Frankenia L. 15 0.96 Sarcocornia A. J. Scott 12 0.77
Salicornia L. 15 0.96 Zostera L. 12 0.77
Salsola L. 15 0.90 Eucalyptus L. Her 11 0.71
Juncus L. 14 0.90 Carex L. 10 0.64
Plantago L. 14 0.90 Rhizophora 10 0.64
Scirpus L. 14 0.90 Anabasis L. 9 0.58
Zygophyllum L. 14 0.90 Casuarina 9 0.58

cific diversity of the С4 plants (36 species) in the flora Table 3. Frequency of occurrence of halophyte species by
life forms
of the Central Karakum, 22 of them are dominant,
while only 15 are dominant out of the 174 С3 species. Number
Life form %
of species
The study of the latitudinal distribution of the С4
halophytes and their confinement to certain ecologi- НР, perennial grasses 403 25.87
cal conditions helped understand in general terms the SH, shrubs 232 14.89
adaptive importance of various mechanisms of СО2 А, annuals 181 11.62
fixation. Using detailed quantitative analysis of the
geographical distribution of the С4 plants in North and CHN, nanochamaephytes 61 3.92
(shrubs not higher than 0.25 m)
Central America, Australia, and Eurasia, a certain
quantitative relation between several climate indica- CH, chamaephytes (small shrubs 125 8.02
tors and the abundance of С4 species was revealed. The less than 0.5 m in height)
high correlation coefficients (r = 0.94–0.97) among AP, annual plants 46 2.95
these characters allow us to conclude about the uni- AQ, aquatics 27 1.73
versality of this relation regardless of the systemic
affiliation of plants. Naturally, the zones of dissemina- G, geophytes 8 0.51
tion of plants with different types of photosynthesis do T, trees 289 18.55
not have clear-cut climatic boundaries because soil VP, climbing perennials 7 0.45
(edaphic) conditions also have a substantial effect.
These two main factors (climate and edaphic condi- V, climbing plants 1 0.06
tions) predetermine the strategy of plant adaptation to F, ferns 4 0.26
arid conditions, which is expressed in the change in SU, succulents 8 0.51
the ratio of plants with various photosynthesis types in
CSH, perennial small shrubs 4 0.26
arid ecosystems in the direction С3 → С4 → САМ
[12]. The С4 species dominate in regions with a hot cli- Р, parasites 2 0.13
mate (not necessarily dry); they can be found in damp SG, seagrasses 37 2.37
and shadowed places, but they do not dominate there. SAQ, seagrasses 4 0.26
In superarid desert conditions, С3 species are practi- (prevail at depths of 0.2–1.5 m)
cally absent and the vegetation is represented by С4 PG, perennial grasses 115 7.38
and САМ species. The САМ species are able to fix
СО2 at night and spend moisture even more econom- Loss 4 0.26
ically than the С4 species [12]. Total 1558 100.00

HERALD OF THE RUSSIAN ACADEMY OF SCIENCES Vol. 87 No. 1 2017

6 SHAMSUTDINOV et al.
Table 4. Annual average yield of halophyte dry matter with
seawater irrigation in Puerto Penasco, 1990–1992
Dry matter yield,
Number t/ha
Halophyte species
of samples
Average SE
Batis maritime 8 33.9 0.99
Atriplex linearis 5 24.2 1.23
Salicornia bigelovii 22 22.4 0.70
(1st year)
Salicornia bigelovii 9 17.7 1.32
(2nd year)
Suaeda esteroa 9 17.2 1.12
Sesuvium portulacustrum 9 16.7 2.00
In addition to climatic factors, the ratio of С3 to С4
species in ecosystems depends on edaphic conditions
(the soil water regime, mechanical composition,
aquatic–physical properties, and salinity level) and
light provision. Within one climatic region, the ceno-
tic role of the С4 plants increases as the soil conditions
deteriorate. According to data in [12, 13], the number
of С4 species in the series “dune sands–fixed sands–
takyr-like soils–solonchaks” in the Karakum Desert
has increased from 16.5 to 70%. This series is also
noted for the deterioration of soil conditions; water
deficit increases; degradation of the agrophysical
properties of the soil; and increase in salt concentra-
tions. In addition, a certain ratio of С3 to С4 species
corresponds to a certain interval on the “scale of
extremeness.” Thus, the definition of a photosynthesis
type makes it possible to assess approximately the cli-
matic and edaphic capabilities of a plant.
Plants with different photosynthesis types are
divided into sufficiently contrastive biochemical and
structural groups, differentiating them by the
edaphic–climatic confinement. According to data in
[17, 18], the С4 grasses and С4 dicotyledons on the ter-
ritory of North America are climatically distributed as
follows: the northern boundary of the С4 grasses is
predetermined by low temperatures, and the distribu-
tion of the С4 dicotyledons mainly depends on the pre-
cipitation regime. The distribution of grasses and
dicotyledons with the С4 photosynthesis depending on
edaphic conditions in various ecosystems of Southern
Tajikistan was demonstrated by P’yankov and
Mokronosov [13]: the С4 grasses dominate on non-
saline, sufficiently moist soils, but, as the water and
salt stresses increase, their share decreases sharply,
and the number of С4 dicotyledons increases.
The further ecological differentiation of the С4
plants among monocotyledons and dicotyledons may
be related to the presence of contrastive structural–
HERALD OF THE RUSSIAN ACADEMY OF SCIENCES
functional types [19]. The С4 plants are divided into
three groups by the way in which dicarboxylic acids
decarboxylate: NADP-malic enzymic, NAD-malic
enzymic, and PEP-carboxykinasic. In addition, the
anatomical diversity of С4 plants prompted the cre-
ation of structural classifications in the grass and
goosefoot families. The special attention in these two
families is explained by the fact that they concentrate
about 80% of the known С4 species. In the Poaceae
family, panicoid, aristidoid, and chloridoid structural
types of the С4 syndrome are distinguished, and in the
Chenopodiaceae family, salsoloid, chohioid, atriple-
coid, and suaedioid types are distinguished. Structural
groups in combination with biochemical ones form
unique structural–functional types of the С4 syn-
drome, association with which predetermines the eco-
logical properties of plants [12].
If we speak about goosefoot representatives, in rel-
atively favorable soil conditions, mainly malate species
with the chohioid and salsoloid structural types are
found. Extreme habitats (takyrs, solonchaks) are pop-
ulated exclusively by aspartate species with the salso-
loid, suaedioid, and atriplecoid structural types of the
assimilating organs. Global and domestic experience
shows that halophytes represent a major biological
tool of utilizing saltwater with the simultaneous pro-
duction of feeds and medicinal and oil raw materials.
Halophytes in crop production. Over 150 halophyte
species in the world’s flora are of feed value. In Aus-
tralia, Mexico, Israel, the United States, and Russia,
50 promising species of halophytic shrubs, subshrubs,
and grasses—saxauls, cypresses, the genus Climacop-
tera, seepweeds, etc.—have been selected [1, 20].
The characteristic features of halophytes as forage
plants are their sufficiently high nutritive value; stable
forage equilibrium by seasons, especially in fall and
winter; and a full value of protein (a high content of
essential amino acids). These features allow using
halophytes as a fattening feed for sheep, goats, and
camels in arid zones [2, 20].
Growing a number of halophytes for feed in single-
crop plantings and mixtures when irrigated by seawater
can furnish a crop yield equal to that of alfalfa irrigated
by freshwater [1]. Acknowledged as most promising
are the species of the genus Atriplex L. Great attention
has been paid to the domestication of these species in
Australia, Israel, the United States, and India. Atriplex
species are used as forage, oil, and decorative plants.
Over the past 25 years, the associates of Arizona
State University have conducted experiments in irri-
gating halophytes with seawater in the following
places: Puerto Penasco, Kino Bay (Mexico, the upper
Gulf of California); the Pacific coast of Baja (Califor-
nia); the Gulf of Oman (UAE); Hurghada (Egypt,
northern territories closer to the Red Sea); and Abu
Dhabi, Dubai, and Kuwait City (the Arabian Gulf).
Table 4 shows data on the average crop yield of halo-
phytes in 1990–1992 in Mexico. Apparently, if irri-
Vol. 87 No. 1 2017
 HALOPHYTES: ECOLOGICAL FEATURES 7

Table 5. Growth, development, and productivity of halophytes in the conditions of a collection nursery upon irrigation with
salt water from underground sources in Central Kyzyl Kum, 1989
Feed mass yield,
Plant stand, Plant t/ha
Sample Origin
thou. ha height, cm
green dry
Climacoptera crassa
К-7 Karakalpakstan, Berunii district, urochishche Altynsai 203.3 ± 2.6 63.3 ± 2.2 114.4 21.1
К-20 Uzbekistan, Bukhara oblast, Kanimekh district, 173.3 ± 2.5 75.3 ± 0.9 96.6 ± 23.8 17.8
sovkhoz 40 Years of October
К-19 Uzbekistan, Xorazm oblast, Khiva district, 166.6 ± 4.1 46.3 ± 1.2 63.3 11.7
Komsomol’skoe Lake
Salsola turkestanica
К-4471 Kazakhstan, Kzylorda oblast, Zhanakorgan district, 116.0 ± 0.98 89.6 ± 2.16 62.2 11.5
vicinity of the village of Akkum
Kochia scoparia
Uzbekistan, vicinity of Samarkand 188.3 ± 10.4 116.6 ± 1.38 – 17.3
Bassia hyssopifolia
Uzbekistan, Bukhara oblast, Kanimekh district, 24.1 ± 1.5 136.4 ± 3.9 64.8 16.2
sovkhoz 40 Years of October

gated with seawater, it would relatively easily be possi-
ble to obtain 17–34 t/ha of dry substance of the phyto-
mass.
The studies show that halophytes irrigated with
seawater are promising for the development of sea and
oceanic coasts [1, 2, 4, 5]. In particular, they can be
used in the system of commercial fish farms, for exam-
ple, for growing shrimp and langoustes. Mexico has
already developed a target technology of growing
halophytes irrigated with seawater and introduced in
fish livestock farms [4].
A series of studies on growing forage halophytes
under saline water irrigation was conducted in the
Central Kyzylkum (2500 mg/L). The soils of the
experimental plot are sandy desert, of a sabulous
mechanical composition; the annual precipitation is
80–130 mm; the annual average air temperature is
15°С; the average January temperature is –1°С; that
of July is +27°С; the duration of the vegetation period
(days with temperatures above 10°С) is 211–220 days;
and the sum of active temperatures is 4000–4500°С.
The data that characterize the growth, develop-
ment, and productivity of halophyte species and their
samples, collected in various natural-ecological con-
ditions of arid zones in Central Asia and irrigated with
ground saline water, show positive dynamics (Table 5).
The annual halophyte Climacoptera crassa has an
exceptionally high intraspecific diversity by the char-
acter of production. Among the forms studied,
sample K-7, gathered in urochishche Altynsai,
Karakalpakstan, accumulates the largest feed mass,
21.0 t/ha; samples K-20, K-11, and K-4345 form
12.9–17.8 t/ha of dry matter. At the same time, Clima-
coptera samples with a low productivity were found
(2.2–4.5 t/ha of dry matter).
An experiment of growing Kochia scoparia (L.)
Schrad. was conducted in the conditions of the Low
Volga region on secondarily salinized soils at the
Astrakhan test-melioration station (15 km from Astra-
khan). Fourteen samples of Kochia scoparia, gathered
in Qashqadaryo (1 sample), Bukhara (1), and Samar-
kand (3) oblasts of Uzbekistan; Chardzhou oblast of
Turkmenistan (2); and Volgograd (4) and Astrakhan
(3) oblasts of Russia were tested. The seeds were
planted into plowed fields under the conditions of a
collection nursery, the seeding depth being 0.5–1 cm.
The Kochia scoparia seedlings appeared in the first ten
days of March. It was established that the duration of
the vegetation period of various samples was 199–
210 days. The comparative analysis of passing through
phenological phases and plant growth dynamics
showed vividly the great difference between the Cen-
tral Asian and Low Volga Kochia scoparia samples.
The Central Asian samples were characterized by con-
siderable tallness and foliage compared to the Astra-
khan and Volgograd samples. The tested samples also
differed in the size of the forming feed mass and the
number of seeds: the range of yield fluctuations was
7.7–14.1 t/ha of dry mass and 0.6–1.3 t/ha of seeds
(Table 6). The most productive were samples K-301
(Guzar district, Uzbekistan), K-303 (Samarkand
oblast, Uzbekistan), and K-345 (Chardzhou, Turk-
menistan), which were able to yield 12.3–14.1 t/ha of

HERALD OF THE RUSSIAN ACADEMY OF SCIENCES Vol. 87 No. 1 2017

8 SHAMSUTDINOV et al.

Table 6. Yield indices of Kochia scoparia (L.) Schrad. samples of various ecological–geographical origins on saline soils of
the Lower Volga by test years. The collection nursery, Astrakhan test-melioration station, Astrakhan oblast
Samples by Plant height, Green mass, Dry Seeds,
Sample-picking location
VIR catalog cm t/ha mass, t/ha t/ha

К-301 Uzbekistan, Qashqadaryo oblast (1) 130.9 21.10 12.65 0.95

К-302 Uzbekistan, Bukhara oblast (2) 134.5 21.25 10.20 0.70
К-303 Uzbekistan, Samarkand oblast (3) 136.4 23.85 11.65 1.00
К-304 Turkmenistan, Chardzhou oblast (4) 143.5 18.95 8.00 0.70
К-305 Turkmenistan, Chardzhou oblast (5) 145.0 22.26 11.05 0.80
К-345 Volgograd oblast (6) 153.4 24.95 12.00 0.95
К-350 Volgograd oblast, Tsaritsyno village (7) 152.5 21.95 10.00 0.75
К-370 Volgograd oblast, Svetlyi Yar (8) 138.9 17.50 7.90 0.70
К-371 Volgograd oblast (9) 146.7 17.25 7.95 0.65
К-378 Astrakhan oblast, Chernyi Yar (10) 148.8 18.65 10.35 0.70
К-381 Astrakhan oblast, Solenoe Zaimishche village (11) 150.8 24.20 11.10 0.85
К-391 Astrakhan oblast, Chernyi Yar (12) 146.3 19.15 9.00 0.85
(1) Light desert soils, mean chloride–sulfate salinization; (2) desert sand soils, heavy chloride–sulfate salinization; (3) light desert soils,
clay-loam soil; (4) gray-brown light loams, heavy salinization; (5) gray-brown soil, close stand of ground water (0.8–1.5 m), heavy sali-
nization; (6) light chestnut soils, loamy sands, medium saline; (7) light chestnut soil, subsaline; (8) chestnut alkaline soils; (9) vicinity of
airport, light chestnut soils; (10) light chestnut alkaline and saline soils, loamy; (11) light chestnut alkaline soil, loamy; (12) abandoned
rice checks, alkaline and saline soil.

dry feed mass and 1.1–1.2 t/ha of seeds. These samples
were characterized by a higher foliage: the leaf and
seed content in the feed mass was 55–56%.
Table 7 shows the results of studies on various eco-
logical–geographical Kochia scoparia samples irri-
gated with the Caspian seawater on Mangyshlak Pen-
insula. The samples developed normally, reaching a
height of 46–136.4 cm by the time of seed maturation.
When irrigated with seawater, Kochia scoparia formed
6–13.2 t/ha of dry matter. The most productive turned
out to be samples K-5, K-35, K-101, and K-102.
Alongside Kochia scoparia, one sample of halophyte
annuals was tested, Suaeda arcuata and Kochia
iranica. There species also turned out to be very prom-
ising for the conditions of the Mangyshlak Desert,
forming 13.6 and 28.1 t/ha of dry matter, respectively.
Oil halophytes. The importance of halophytes as
potential oil crops is great. The halophyte annual Sa-
licornia bigelovii has by now been domesticated in the
United States, Mexico, Saudi Arabia, and Egypt; a
variety of this halophyte has been created, SOS-10. It
is an annual plant, practically leafless, succulent; it
grows on salinized marshes and clay loams and propa-
gates by seeds. Salicornia SOS-10, when irrigated with
seawater, forms 20 t/ha of dry matter, 2 t/ha of seeds
with 30% oil content and ensures the production of
600 kg of oil from 1 ha. The total price per 1 ha of the
cultivated crop is $600–$650, the cost of production
of 1 t of the seeds being $300–$350 [21]. The oil
extracted from the seeds and further refined by coarse
and fine filtration is used in food production and
industry. The cake and meal from Salicornia seeds
contain up to 40% of protein and are suitable to use as
cattle feed [2, 4].
The wild-growing shrub jojoba (Simmondsia chin-
ensis C.K. Schneider) has been domesticated in the
United States, Mexico, and Israel. Its most valuable
property is its high resistance to salinization. Jojoba
grows in areas where the level of saline groundwater is
1.8 m deep. In Israel, commercial plantations of this
plant are laid down on the coast of the Dead Sea. The
oil content in the seeds is about 50%, and the protein
content is up to 35%. The practical value of jojoba
depends on the unique quality of the oil produced
from its seeds. Lubricants are also made from jojoba;
they preserve viscosity at high pressures and low or
high temperatures, making them usable in high-speed
machinery. The oil’s purity and resistance to rancidity
make it usable as a high-quality base in pharmacology
and cosmetics. Large enterprises that produce lubri-
cants, research institutes, and cosmetics companies of
Europe, Mexico, and the United States purchase the
oil at rather high prices, from $3000 to $20 000 per 1 t.
Of great practical interest is research into the irriga-
tion of jojoba with seawater (the United States and
Israel). In Israel, at least 25% of the area planted with
jojoba is irrigated with waste and saline water. The
plant reacts positively to seawater irrigation at a 35–
40 g/L mineralization [2, 21].

HERALD OF THE RUSSIAN ACADEMY OF SCIENCES Vol. 87 No. 1 2017

 HALOPHYTES: ECOLOGICAL FEATURES 9

Data accumulated abroad on jojoba domestication Table 7. Productivity of Kochia scoparia samples irrigated by
show that it belongs to the most promising plants for Caspian seawater (Mangyshlak, Kazakhstan)
arid regions. Especially great is its practical value for Feed mass
the development and economic recovery of countries Samples Plant stand, Plant yield, t/ha
and regions with scarce natural resources. by catalog thou./ha height, cm
Note that in many cases the nutritional value of oil green dry
from halophyte seeds is higher than from traditional
Kochia scoparia 68.9 71.7 18.0 8.0
oil crops. For further research in this field, the follow-
ing halophytes are recommended: Distichlis spp., Vou- К-37
vea spp., Allentrolfea accidentalis O. Kuntze, and К-57 36.7 101.6 28.1 11.3
Suaeda forreyana Hook. & Aen. К-35 81.3 80.9 33.0 12.0
Medicinal halophytes. The licorice genus Glycyr- К-51 26.7 88.3 25.0 9.7
rhiza L. includes 13 species, 7 of which grow in the CIS
countries and 3, in the northwest of Russia. However, К-3 32.4 96.8 18.0 7.8
only the mesohalophytes common licorice, G. glabra L., К-101 15.6 134.7 36.1 13.2
and Ural licorice, G. uralensis Fisch., are of commer-
К-92 14.6 148.7 15.2 6.0
cial value as medicinal, feed, and biomeliorative
crops. К-93 24.1 136.4 41.5 16.2
The licorice root is a valuable pharmacological, Suaeda arcuata 15.3 113.6 54.4 13.6
food, and technical raw material [22]. It is included in К-58
the pharmacopoeias of over 30 countries, holding first Lochia iranica 29.7 205.9 112.0 28.1
place among medicinal plants in procurement vol-
umes. Licorice has become very popular primarily
thanks to the glycyrrhizic acid contained in it, the con- A number of commercial enterprises in the United
tent of which in underground organs ranges from 3 to States and other countries specialize on marketing
20%. Preparations for the treatment of allergies, bron- ornamental plants and their seeds [1]. In the south of
chial asthma, cold relief, and expectorants have been Israel, salt-resistant plants have been used as greenery
made and are widely used on the basis of glycyrrhizic for two decades [5]. The most valuable species selected
acid. Moreover, on the basis of flavonoids, which are and disseminated by the Institute of Applied Research in
also contained in the licorice root, preparations were Israel include trees (Melaleuca halmaturorum F. Muell.
developed for the treatment of gastric and duodenal ex Miq., Tamarix L., Conocarpus erectus L.), shrubs
ulcers, hyperacid gastritis, and other illnesses that (Borrichea spp., Clerodendron inerme R. Br., Maireana
require wound-healing and hemostatic actions [22]. Moq., Seaevala spp.), and ornamental undersized and
Licorice hay has satisfactory feed properties and is trailing plants (Crithmum maritimum L., Gallnia spp.,
used in agriculture as a silage (mixed with corn) and Drosanthemum spp., Halimus portilucoides Wallr.,
hay plant. Licorice hay laid in during the fruiting stage Limonium spp., Lippia nodiflora Michx. Fl., Sesuvium
is close to good-quality grass hay nutritionally. Com- spp.) [5, 25].
mon licorice produces 10–12.4 t/ha of hay [23]. At present, Israel is one of the largest exporters of
Halophytes as ornamental plants. The use of land- ornamental halophytic plants. The annual profit from
scape halophytes to replace glycophytes or in areas this export is over $90 mln [1, 2, 21].
where glycophytes cannot grow has a great aesthetic Halophytes as energy inputs. Halophytes are used
and practical potential [1, 20]. The coasts of oceans as wood fuel. The United States has developed a tech-
and seas and other regions of the world where soils are nology of phytomass bales from halophytes. Planta-
salinized and water resources are limited, have inex- tions of halophytic plants are energy-producing
haustible possibilities for raising ornamental halo- renewable biological resources and simultaneously
phytes [20]. energy storages.
The formation of landscapes using halophytes is a Some halophytes, including Casuarina Miq., Tam-
means of controlling extreme climate conditions; it arix L., and Haloxylon Bunge., and some Lagonychium
mitigates negative impacts on the environment and is Bieb. species, are recommended as energy inputs in
often more economical than measures such as light, saltwater irrigation. To produce biomass for energy
wind, and noise protective structures [24]. purposes, Atriplex canescens (Pursch.) Nutt., Bigelowia
Hot and arid regions with limited resources of DC., Sarcobatus vertmiculatus (Hook.) Toor., Arte-
potable water in countries of the Middle East, Central misia tridentate Nutt., and Haloxylon ammodenron
Asia, and Africa could profit from promoting a pro- (C.A. Mey.) Bunge are recommended for irrigation-
gram of ornamental halophytic crop production. To free growing [2, 21].
varying degrees, 240 halophyte species of various life Each acre of halophytic annuals planted produces
forms are of ornamental value [1, 25]. energy equal to 1250 gallons of gasoline, and 100 acres

HERALD OF THE RUSSIAN ACADEMY OF SCIENCES Vol. 87 No. 1 2017

10 SHAMSUTDINOV et al.
of a ten-year-old plantation of a halophyte tree,
the black saxaul, produce energy equivalent to
625 000 gallons [1].
Halophytes as biomeliorants. Halophytes have hab-
itat-forming and habitat-optimizing functions and
consequently create an ameliorative effect on salinized
soils. Thanks to the effective shadowing of the soil sur-
face by the top mass, pumping functions, and biologi-
cal-drainage functions, halophyte plantations ensure a
sharp decrease in physical evaporation, a reduced
groundwater level, salt carryover by the top mass, and
soil desalinization [20, 23]. The fresh organic matter
supplied by plants helps improve the physicochemical
properties and biological activity of the soil, as well as
change the рН, electrical conductivity, and hydraulic
conductivity.
Domestic experience shows that the main principle
of developing meliorative crop rotation consists of
using halophytes during the first years with further
transfer to mixed plantings of a halophyte with a feed
crop and a gradual, as the soil desalinizes, increase in
the area for the feed crop. When the soil has been
desalinized completely, a pure feed crop is planted
[20]. The biological approach to soil desalinization is
recommended for medium saline and highly saline
medium-loam soils, when the degree of chloride sali-
nization does not exceed 0.6%.
The research tasks for the next few years boil down
to the need to expand the mobilization of global plant
resources of halophytes at the specific and ecotypic
levels, create their genetic pool, study comprehen-
sively and assess the habitat-forming functions of
halophytic plants, and develop adaptive methods of
selection and breeding of ecologically differentiated
and economically specialized varieties, as well as
design the technologies of cultivation and biological
melioration of degraded agrolandscapes in arid
regions of Russia and Central Asia.
ACKNOWLEDGMENTS
This work was supported by the Russian Founda-
tion for Basic Research, project no. 15-05-08025,
“Halophytes: Global Genetic Resources, Biological,
Ecological–Physiological, and Production Charac-
teristics and Prospects for the Practical Use of Exper-
imental Botany.”
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Translated by B. Alekseev
Vol. 87 No. 1 2017