Physiology of Domesticated Alliums: Onions, Garlic, Leek, and Minor Crops
R Kamenetsky, Agricultural Research Organization, Bet Dagan, Israel
HD Rabinowitch, The Hebrew University of Jerusalem, Jerusalem, Israel
2017 Elsevier Ltd. All rights reserved.
Introduction
For millennia, members of the genus Allium L. (Amaryllida-
ceae) have been cultivated worldwide for sustenance, avor,
and medicinal purposes, and recently also as ornamentals. Of
the ca 800 Allium species, more than 50 are cultivated in ca
175 countries, from the tropics to the temperate zone where
they usually store well and are consumed as fresh and pro-
cessed foods.
The most common Allium food crops are garlic, leek,
shallot, chives, Welsh onion, and bulb onion (van der Meer,
1997; Table 1). The latter are the second leading world vege-
table preceded only by tomatoes.
Genetic Resources, Variability, and Improvement
The center of Allium diversity stretches between Central and
Eastern Asia to the Mediterranean Basin, where, most likely,
garlic and onion were rst domesticated. Western North America
is considered a second smaller center of diversity (Meerow, 2012;
Friesen et al., 2006; Li et al., 2010).
Bulb onion (Allium cepa L.) and shallot (Allium cepa Aggre-
gatum Group) are not known in nature, but their closest wild
relatives, Allium oschaninii O. Fedtsch. and Allium vavilovii M.
Popov et Vved., grow wild in Central Asia. The domesticated
bulb onions fall into two main horticultural groups. All culti-
vars of the seed propagated Common Onion group form large
single bulbs, and following maturation and/or owering, their
axillary meristems develop into dormant lateral buds. Onion
cultivars vary markedly in many physiological and horticultural
traits, for example, in response to photoperiod, to temperatures
during storage and growth, in dry matter content, in avor,
pungency, and doubling in shape, in the number and adhesion
of tunics, and in skin and esh color (Brewster, 2008).
Members of the onion Aggregatum Group produce small
dividing bulbs. Independent of the apical domes differentia-
tion to the reproductive phase, new lateral meristems develop
at the axils of the leaves, grow, and form clusters of bulblets.
Their propagation is mainly vegetative but recently production
of shallot from seed has become common (Rabinowitch and
Kamenetsky, 2002).
Table 1 Production of Allium crops: acreage and tonnage
in 2012 FAO (http://faostat3.fao.org/faostat-gateway)
Production
Crop Tonnes Area, ha
Dry onions 82 851 732 4 203 648
Dry garlic 24 836 877 1 465 772
Green onions, shallots 4 342 135 239 594
Leek, other vegetable alliums 2 165 667 132 648
Encyclopedia of Applied Plant Sciences, 2nd edition, Volume 3 http://dx.doi.org/10.1016/B978-0-12-394807-6.00064-2
In addition to matured plants, leaves of young alliums of
both groups are consumed prior to, and shortly after bulbing
begins as salad onion (scallions), without or with developing
bulbs, for their delicate allium avor.
Garlic (Allium sativum L.) is known solely in culture and due
to complete sterility it is only propagated vegetatively from
cloves. In the absence of sexual reproduction, mutations,
whether at random in intact plants or when propagated in
tissue culture (somaclonal variation) are the only source of
variation. Nevertheless, garlic clones vary markedly in bulbing,
bolting bulb size, numbers of whorls and cloves, skin color, dry
weight, and pungency. They also differ in response to temper-
atures and photoperiod, with the consequent adaption to
a variety of ecological zones. Sterility, however, markedly holds
back improvements by classical breeding of economically
important traits, including pest and disease resistance, yield,
and quality (Kamenetsky, 2007).
Marked variation among garlic clones with respect to ow-
ering ability and the ratio between owers and topsets in the
umbel suggested that garlic undergoes a process of transition
from sexual to asexual reproduction. Evidently, the ancestral
garlic owered, had normal meiosis, and produced seed in
the long-scape multiower umbel. Compared with modern
cultivars, the ancestral garlic from the high plateaus of Central
Asia probably had greater tolerance to extreme temperatures
and matured later than the domesticated plants did. The devel-
opment and growth of owering scapes consumes energy at the
expense of the storage organs. It is therefore theorized that
selections for early maturation of big garlic bulbs deprive the
developing scapes from essential energy and nutritional
supplies, thus promoting sterility of the domesticated plants.
Hence, expeditions went to the primary center of garlic
evolution for collection of owering plants. Forcing of these
accessions and rigorous selections in Japan, the United States,
and Israel thus resulted in lush owering and improved seed
production. Additionally, physiological studies and environ-
ment manipulations led to blooming of fertile inorescences
in many genotypes thus indicating that the genetic makeup
that controls blooming and fertility is by and large intact.
Seed-derived garlic progenies vary markedly in morpholog-
ical, physiological, economical, and horticultural important
traits, such as tolerance to rust, Puccinia allii. This variability
provides a high potential for improvement by breeding and
selection among sexually derived progenies with the conse-
quent development of superior cultivars and introduction of
new traits currently uncommon in commercial clones.
Allium ampeloprasum L., the common ancestor of leek, kur-
rat, pearl onion, and great-headed garlic (elephant garlic),
grows wild all around the Mediterranean basin from Portugal
to western Iran. Of these, only great-headed garlic forms bulbs
and cloves similar to garlic, but size, number of cloves per bulb,
and avor are markedly different.
Leaves of the seed propagated nonbulbing perennial
Japanese bunching onion (also known as Welsh onion: Allium
255
256 Horticulture Production and Quality j Physiology of Domesticated Alliums: Onions, Garlic, Leek, and Minor Crops
stulosum L.) and the bulbous Rakkyo (Allium chinense G. Don)
are popular in Eastern Asia. The former is widely cultivated
around the world as a substitute for scallions. In China, Japan,
Korea, Taiwan, and Indonesia, where most of the world bunch-
ing onion production is made, it is a very important vegetable,
ranking among top 10, and is marketed all year round.
Chives (Allium schoenoprasum L.) and Chinese chives (Allium
tuberosum Rottl.) are grown mainly in Europe and Asia for their
allium-avored leaves. They do not form bulbs, but produce
nonedible storage rhizomes, used for vegetative propagation.
Standard propagation of chives, however, is from true seed.
Numerous minor crops with typical alliums avor are grown
in culture, for example, French gray shallot (A. oschaninii), Allium
cepa viviparum (also known as tree onion, top onion, and
Egyptian onion (Allium
proliferum)), or collected in the wild,
for example, Chinese or Japanese garlic (Allium macrostemon
Bunge), Naples garlic (Allium neapolitanum Cyr.), ramsons
(Allium ursinum L.), long-rooted garlic (Allium victorialis L.),
Canada garlic (Allium canadense L.), ramp (Allium tricoccum
Ait.), Allium motor Kamelin & Levichev., Allium pskemense B.
Fedtsch., Allium galathum Kar. et Kir., and Allium altaicum Pall.
In terms of breeding and improvement, the edible alliums
fall into four major groups: the fertile bulb onion, leek, Japa-
nese bunching onion, and chives, where classical breeding
brings about new improved varieties. On the other hand, garlic
and elephant garlic are sterile with the consequent little room
for genetic and breeding works. Chinese chives displays an
intermediate position as it produces apomictic seeds geneti-
cally identical to the mother plant, yet cross pollination is
achievable and therefore conventional breeding is possible.
The key agents supporting cross-pollination in alliums are
insects, with the consequent indenite conservation of some
degree of heterozygosity. Additionally, cross-pollination leads
to incorporated variation and exhibition of heterosis. Hence,
most modern cultivars of onion, Japanese bunching onion,
leek, and some modern shallot cultivars are hybrids, and seed
production depends on stable inherited male sterility.
All alliums, both diploids (e.g., bulb onion: 2n 2x 16)
and tetraploids (e.g., leek: 2n 4x 32) possess some of the
largest genomes in the plant kingdom, probably due to intra-
chromosomal duplications. The genome sizes range between
10 and 20 Gbp, c.30 times larger than that of rice and similar
to the hexaploid wheat genome (Ricroch et al., 2005). Hence
genome sequencing of Allium species is a challenging task.
Recently, sequencing and mapping of the onion genome has
been initiated by collaborative international efforts. By now,
a comparative genomics database has been developed for
onion, Japanese bunching onion, and Allium roylei Stearn
(McCallum et al., 2012). Transcriptome data is available for
onion (Duangjit et al., 2013) and garlic (Sun et al., 2012;
Kamenetsky et al., 2015). Molecular markers for color, quality
traits, and cytoplasmic male sterility (CMS) are available for
onions (Havey, 2013).
Annual Cycle, Morphology, and Environmental Effects
on Plant Development
Most fertile cultivated alliums are raised from seed and to
a lesser extent from transplants, or sets. The true leaf of bulb
onion consists of a concaved cylindrical, attened blade, and
a sheath attached to the basal plate (squat compressed stem).
Formed in the apical meristem, each new leaf is enveloped by
the sheath of the previous one, thus making for a series of
concentrically arranged round leaf bases that jointly form a false
stem (pseudostem). Following bulb formation in bulb onion,
scales are formed from sheaths and lateral buds. In both garlic
and elephant garlic, these lateral buds develop into cloves.
These cloves do not develop into leaf blades, but serve for
storage of reserves and for propagation. When dormancy
breaks, growth renews from sprouting lateral buds.
Genetic makeup, photoperiod, and both storage and
growth temperatures are the main factors controlling
Alliums bulbing and orogenesis (Mann and Lewis, 1956;
Takagi, 1990; Rabinowitch, 1990; Kamenetsky et al., 2004;
Kamenetsky and Rabinowitch, 2006). The strong differences in
morphological, biochemical, and physiological traits between
and within species probably occur due to a long history of
isolated selections for desired traits, for example, multiple bulbs
in shallot versus large single bulb in onion, or nonbulbing leek
versus aggregate (cluster) of cloves in elephant garlic.
Bulbing
Bulbing of onions and shallots is induced mainly by exposure
to long photoperiod (LD), and red:far-red light ratio plays an
important role. The light signal is sensed by the youngest
emerging leaf; hence its pruning prevents bulbing initiation
or holds up the continuation of the process. Sensitivity to LD
is almost (but not entirely) independent of plant age or
mass. It ranges from about 10.5 light hours (e.g., cv. Early
Beit Alpha) to about 16 light hours for the late maturing types
common in high latitudes. Based on minimum light hours
requirements, allium crops are classied as short, intermediate,
or LD cultivars.
Many other environmental factors affect bulbing and bulb
size, including temperatures, especially night temperatures, irri-
gation, mineral nutrition, weed competition, and sand.
Preplanting storage affects bulbing. Following storage at
2830  C, onion and shallot bulb later than those grown
from sets stored at <20  C. Similarly, following storage at
210  C, plants raised from cloves of the Israeli garlic cv Shani
bulb earlier and bear markedly lower yields than those raised
after clove storage at 30  C. Garlic varieties from cooler regions
however, respond differently. Thus, temperatures below 15  C
are required for bulb induction in Japanese garlic cultivars.
In bulbing alliums, even late interruption in LD exposure
results in arrest of the bulbing process and in sprouting of axil-
lary buds. The latter sprouting consumes energy stored in the
scales, thus the developing main bulb shrinks.
The size of mature bulbs depends on the total number of
existing leaves and leaf buds when bulbing initiates, on growth
conditions and the amount of assimilates stored in the leaf
blades and sheaths prior to the commence of bulbing, and
on further assimilation during the bulbing process. Longer
growth and assimilation period may result in larger bulbs
and higher yields.
When bulbing begins, leaf emergence and cell division stop,
and leaf bases and young buds become strong sinks; both leaf
sheaths and leaf buds turn into thick false and true scales,
 Horticulture Production and Quality j Physiology of Domesticated Alliums: Onions, Garlic, Leek, and Minor Crops 257

respectively. Concomitantly, their cells swell to accommodate
the basipetal ow of sugars, for energy storage and supply
needed during dormancy and renewed growth. The levels of
growth retardants increases, apical cell division stops, and the
oldest sheaths turn into protective tunics. The cessation of
new leaf emergence leads to the formation of empty space
inside the pseudostem. This mechanical weakening together
with the turgor dwindling due to the growing ow of assimi-
lates downward leads to foliage collapse. Yet, photosynthesis
in the green blades continues thus contributing to the bulb
growth and yield. Gradually blades get dry and die back in
the sequential order of development, i.e., older leaves rst.
When all leaves die out, the mature bulb enters into a phase
of physiological rest/dormancy.
After harvest, there is a temperature-dependent breakdown
of the growth retardants, a concomitant weakening of the
bulbs rest/dormancy and the consequent sprouting.
Floral Development
With a few exceptions, vernalization (prolonged exposure to
low temperatures) is required for oral induction in bulb
onion (optimum 9  C, range 017  C); shallot (range
510  C); leek (optimum 5  C, range 018  C); and garlic
(range 2 to 10  C). Floral initiation occurs only in postjuve-
nile plants or resting bulbs after having a minimum of 5, 6,
or 714 leaf primordia and leaves in leek, shallot, and in
bulb onion, respectively. This sensitivity to the external signals
increases with physiological age. In Chinese chives, leek, and
rakkyo, LD signals are required for oral induction.
In contrast, eld or storage temperatures of 2830  C
inhibit oral initiation in onion and cause a owering delay
during the subsequent growing season or great reduction in
owering. Yet, in leek, high temperatures do not lead to abor-
tion of already developed umbels.
Most garlic cultivars and some shallot genotypes do not
produce reproductive organs, possibly due to a long history
of selection against this trait.
In onions, molecular physiology studies show that ower-
ing and bulbing inductions are closely connected and are regu-
lated by similar genetic mechanisms. Indeed, several members
of the Flowering Locus T (FT) family of genes are involved in
both processes (Lee et al., 2013). Flowering promotion by
vernalization correlates well with the upregulation of AcFT2,
whereas bulb formation is regulated by two antagonistic
FT-like genes (Figure 1). Under short photoperiod, the AcFT4
gene products prevent the upregulation of AcFT1 and bulbing.
In contrast, exposure to LD induces the downregulation of

Figure 1 Under short day/short photoperiod, early bulbing in year 1 of spring planted onion seedlings (left) is initially inhibited due to AcFT4 expres-
sion. Once the critical day-length is reached, AcFT4 is downregulated with the consequent bulb induction by the upregulation of AcFT1. Overwintering
and cold exposure (vernalization) leads to AcFT2 upregulation with the consequent induction of owering in the second year. Flowering and seed
setting occur in the following spring/summer. Lee, R., Baldwin, S., Kenel, F., McCallum, J., Macknight, R., 2013. FLOWERING LOCUS T genes control
onion bulb formation and owering. Nat. Commun. 4 (2884), 19. http://dx.doi.org/10.1038/ncomms3884, with permission from Nature.
258 Horticulture Production and Quality j Physiology of Domesticated Alliums: Onions, Garlic, Leek, and Minor Crops

AcFT4 and the upregulation of AcFT1, with the respective
consequent promotion of bulbing.
In all studied alliums, sensing the vernalization cues for
owering is possible only in postjuvenile plants. A combina-
tion of endogenous and environmental factors induces transi-
tion from the vegetative to the reproductive phase. Later, the
apical meristem produces a distinctive reproductive architec-
ture: the inorescence initials, followed by differentiation of
individual owers (Figure 2(a)2(d)). In ower buds, two
outer whorls of tepals are formed, while the two inner whorls
consist of reproductive organs, including stamens and gynoe-
cium. Florogenesis proceeds with scape elongation and ino-
rescence differentiation, and ends with anthesis. In a single
ower of onion, shallot, and other Allium species, each perianth
lobe and the subtended stamen arise simultaneously from
a single primordium.
Bulb onion and Japanese bunching onion cultivars vary
both in juvenile age and in cold requirement, hence oral initi-
ation and hybrid seed production require specic manipula-
tions for synchronous owering (nicking) including plants
physiological age and storage temperatures.
Leek has an obvious juvenile phase of about ve visible
leaves. In older plants, both ower induction and development
depend on vernalization. When temperatures increase from 12
to 18  C, a decrease in the number of owering plants and an
increase in the mean number of leaves before ower initiation
are evident. Some genotypes bolt at temperatures between 18
and 24  C, but owers do not form even in plants bearing 45
leaves. Under LD, ower induction is delayed compared with
short day (SD)/short photoperiod conditions.
Garlic clones vary in bolting ability; hence classied as
complete, incomplete, or nonbolting genotypes. Genotypes from
the temperate zone require colder and longer induction for
bolting compared with plants from the subtropics and tropics,
probably because of selection against bolting. Under appro-
priate environmental conditions, plants of the rst two groups
produce scape and oral initials. However, concurrent develop-
ment of the vegetative topsets in the inorescence causes early
abortion of ower buds. In addition to the genetic makeup,
scape elongation in garlic depends on environment, that is,
cold preplanting storage (2 to 9  C), low-eld temperatures,
LD, and excessive mineral nutrition promote scape elongation
in incomplete bolters and, under certain conditions, even in
nonbolters, high storage, or eld temperatures, and SD inhibit
scape elongation in bolting plants. Hence, characterization of
garlic genotypes under a single set of conditions may be irrele-
vant in different regions. This environment-dependent
response poses a signicant doubt on the validity of the phys-
iological descriptors compiled only in one or a few climatic
regions (e.g., gene banks for long-day alliums).
Physiological studies provided a solution to the enigma of
garlic sterility and offered environmental tools for owering
regulation and fertility restoration. The competition by the
simultaneously developing bulb and ower sinks determines
differentially the fate of the elongating stalks, ower develop-
ment, and seed production. In garlic, LD triggers the initial
scape elongation, but long exposure to LD interferes with
ower development by induction of topsets development
(Figures 3(a) and 4). The understanding of oral physiology
facilitated intact orogenesis by sophisticated environmental

Figure 2 Florogenesis in Alliums (original pictures taken from Allium ampeloprasum). (a) Vegetative apical meristem (differentiating to leaf
primordia) turns into reproductive apex. Meristem dome enlarges and begins to produce ower initials. Bar 100 mm. (b) First ower meristems are
visible (arrows). Bar 100 mm. (c) Differentiation of ower buds. Following the formation of six anthers (a), two whorls of perianth lobes (pl)
develop. Gynoecium differentiates at the middle of each ower bud. Bar 100 mm. (d) An uneven nonsynchronous differentiation of owers in
a single inorescence. Flower buds at various stages of development are visible. Bar 1 mm.
 Horticulture Production and Quality j Physiology of Domesticated Alliums: Onions, Garlic, Leek, and Minor Crops 259

Figure 3 Flower and topsets development in bolting garlic (Allium sativum). (a) Numerous topsets in the developing inorescence with the conse-
quent ower abortion. (b) Intact inorescence contains normal owers and produces hundreds of viable seeds.

Figure 4 Schematic presentation of owering, bulbing, and dormancy regulation in bolting garlic (Allium sativum) by environment. Quite similar
environmental signals induce owering and bulbing. The two processes coincide with the consequent strong competition for the same resources.
Manipulation of the factors involved in regulation can easily cause deviation from the main route, for example, thus bulb formation instead of ower-
ing; sprouting of axillary buds instead of dormancy; or topset production instead of owers.

manipulation, both prior to and after planting. Consequently,
viable owers in topset-free umbels were obtained. However,
normal owering cannot be achieved when any of the orogen-
esis developmental stages is retarded, interfered with, or
proceeds in the wrong direction (Figure 4).
Propagation
Vegetative reproduction of garlic, elephant garlic, tree
onion, topsetting onions, walking onions, Egyptian onions
(Allium
proliferum), and gray (Griselle) shallot is exclusively
vegetative. This clonal propagation allows for preservation of
quality traits and xation of heterozygosity/homozygosity
levels. Hence, any phenotypic variation within the population
is credited to environmental effects or to infrequent mutations.
The main drawbacks associated with allium vegetative prop-
agation are as follows: (1) low multiplication rate and high
production cost; (2) need for large storage space; (3) great los-
ses in storage due to pathogenic and physiological injuries; and
the most serious disadvantage is (4) the perpetuation of viral
and other infections due to pathogens and the consequent
260 Horticulture Production and Quality j Physiology of Domesticated Alliums: Onions, Garlic, Leek, and Minor Crops
marked reduction in yield and contamination of production
elds (Salomon, 2002). The latter two adverse downsides
lead to heavy yield losses of up to 50%.
Early in the 1970s, micropropagation in vitro proved advan-
tageous over clove propagation due to the high production
rates of pathogen-free propagules by means of meristem-tip
culture combined with thermotherapy and chemotherapy.
A 4-year cycle, however, is required for the massive production
of these propagules. Hence, these propagules are expensive, yet
plants raised from such a material commonly produces signif-
icant and up to 100% more yield of quality bulbs than those
raised from standard cloves.
Seed propagation is common in onion, leek, Japanese bunch-
ing onion, and chives, as well as in Chinese chives, which
produces apomictic seeds. The two basic seed-production
systems in the biennial alliums are seed-to-seed and the
seed-to-bulb-to seed cycles. The latter allows for rouging off-
types (plants that show undesirable variation from the standard)
thus assuring production of true-to-type seed (Peters, 1990).
Leek seed heads develop slowly, thus often harvested late in
the autumn, as compared with spring or summer seed matura-
tion of bulb onion, shallot, garlic, and bunching onion.
Onion and other allium seeds are short lived. Only proper
growth, harvest, and storage conditions enable long-time
storage. The main good storage practices include low tempera-
tures, low humidity, minimum contamination with storage
pests, and gentle handling.
Chemical Composition, Medicinal, and Neutraceutical
Traits
Allium cells contain odorless, stable nonvolatile amino acids of
the general name S-alk(en)yl cysteine sulfoxides (ACSOs),
probably serving for defense against predators, parasites, and
diseases (Keusgen, 2002; Block, 2010).
Of the various ACSOs, alliin is one of the main constituent
of many alliums. When fresh allium tissues are disrupted
(wounding, crushing, or chewing), the vacuolic enzyme allii-
nase catalyzes the synthesis of a wide range of unstable organo-
sulfur compounds from alliin including the main thiosulnate
allicin. At room temperature, allicin quickly breaks down and
gives rise to the partially stable ajoene, methyl ajoene, and
dithiins. These and other ACSOs contribute to the therapeutic
attributes of fresh alliums.
Alliums avor intensity and quality are affected by both
genetic and environmental factors. Within and between geno-
types, marked differences in avor characteristics and nutraceu-
tics potential arise from genetic variability, differential sulfur
uptake, and its metabolism.
In addition to organic sulfur compounds, alliums serve as
rich sources of avonoids (potent antioxidants) and anticlot-
ting agents. In bulb onion, the major avonol glucosides
include quercetin-3,40 -O-diglucoside (3,40 -Qdg) and quer-
cetin-40 -O-glucoside (40 -Qmg), both comprising >80% of the
total avonol content in this plant, reaching 2.1% of the dry
matter of some red onion bulbs.
The main nonstructural carbohydrates in Allium species
consist of glucose, fructose, and sucrose, together with a
series of oligosaccharides, fructans, and fructosans. Alliums
tissues also contain selenium at up to 70100 mg per 100 g
fresh garlic weight, as well as proteins, pectin, minerals, sapo-
nins, and polyamines. Vitamins, such as ascorbic acid, thia-
mine, riboavin, niacin, pantothenic acid, and vitamin E,
have been reported.
To benet from allium therapeutic qualities, a daily intake
of 50100 g of fresh A. cepa and 2.54 g of A. sativum is recom-
mended. Case-controlled studies have shown that a regular
daily intake of both may reduce the risks of some cancer
diseases, for example, gastrointestinal cancers. It is also sug-
gested that daily intake of A. sativum signicantly lowers the
incidence of atherosclerosis-derived diseases.
Extracts from onion and garlic exhibit a strong antibiotic
activity. However, no formulation of alliums extracts has
proved to be a reliable substitute for modern antibiotics.
A. cepa extracts exhibit antiasthmatic and antidiabetic activities,
while A. sativum showed lipid-lowering effects, antiplatelet
activity, and antiatherosclerotic activities.
Horticulture and Storage
The effects of nutrients and water use efciency on alliums
productivity have been thoroughly studied (Jones and Mann,
1963; Brewster and Rabinowitch, 1990; Brewster, 2008). The
shallow root system requires well-drained light-rich soils. The
plants are susceptible to excessive dryness. When this happens,
feeder roots are injured with the consequent production of
small irregular bulbs.
As for most leafy vegetables, the fresh shoots of leek, scal-
lions, green shallots, chives, and Japanese bunching onion
have only a short life, thus cold storage, high moisture, and
modied atmosphere are benecial. The bulb-forming crops,
onion, shallot, garlic, and elephant garlic have long storage
life, depending on the genotype, storage temperatures, and
air moisture. Storage under ambient conditions requires good
ventilation, but long storage requires refrigeration (02  C).
Extended storage of onion and garlic dormant bulbs is also
plausible at 30  C. On the other hand, storage at 15  C
promotes cell division, differentiation, elongation, and leaf
growth. Preharvest application of sprouting inhibitors (e.g.,
maleic hydrazide) for destruction of axillary buds, with the
consequent extension of storage life, is employed in onion
and garlic, except when sets are used for propagation and/or
bulbs aimed at seed production.
Concluding Remarks
The genus Allium consists of a large and variable group of
economically important species, common all over the
world. Yet, most cultivated alliums lack many important
traits, including resistance to pests and abiotic stress. Strong
gene
environment interactions markedly affect growth,
development of storage organs, owering, and seed produc-
tion, but allium physiology is still far from being fully under-
stood and molecular knowledge of alliums is also rather poor.
Utilization of both classical and novel tools for genetic and
physiology studies, including genome sequencing, develop-
ment of molecular markers for economically important traits,
 Horticulture Production and Quality j Physiology of Domesticated Alliums: Onions, Garlic, Leek, and Minor Crops 261

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