Genet Resour Crop Evol
https://doi.org/10.1007/s10722-018-0641-6
RESEARCH ARTICLE
Maguey (Agave salmiana) infructescence morphology and its
relationship to yield components
Minerva Huerta-Lovera . Cecilia Beatriz Peña-Valdivia . Antonio Garcı́a-Esteva .
Josué Kohashi-Shibata . Huitziméngari Campos-Garcı́a . Juan Rogelio Aguirre-Rivera
Received: 22 October 2017 / Accepted: 16 April 2018
Ó Springer Science+Business Media B.V., part of Springer Nature 2018
Abstract Agaves are long-lived semelparous plants
that produce a high number of seeds, in dehiscent
capsules, on the apical section of a stalk, up to 5 m
long, after 8–25 years. These and other characteristics
such as yield and yield components are scarcely
evaluated in the plants of the Agave genus. The
objective of this study was to quantify the capsules and
seeds yield of A. salmiana Otto ex Salm-Dyck plants
simultaneously maturing and growing in the same
region. Infructescences of three plants simultaneously
growing at San Luis Potosı́, Mexico, were harvested.
On them we evaluated the number of umbels, capsules
and seeds (normal and sterile) and their mass per plant.
The study was developed on a completely randomized
design with each infructescence as an experimental
M. Huerta-Lovera  C. B. Peña-Valdivia (&) 
A. Garcı́a-Esteva  J. Kohashi-Shibata
Postgrado en Botánica, Colegio de Postgraduados,
Carretera México-Texcoco km 36.5, Montecillo,
56230 Texcoco, Mexico, Mexico
e-mail: cecilia@colpos.mx;
cecibetipv@gmail.com
H. Campos-Garcı́a
International Maize and Wheat Improvement Center,
Carretera México-Veracruz Km. 45, El Batán,
C.P. 56237 Texcoco, Mexico, Mexico
J. R. Aguirre-Rivera
Instituto de Investigación de Zonas Desérticas,
Universidad Autónoma de San Luis Potosı́, Altaı́r 200,
Colonia del Llano, 78377 San Luis Potosı́, S.L.P., Mexico
unit. Also, the type distribution of the number and
mass of these yield components along the stalk was
evaluated with the Chi square test for goodness of fit,
the Shapiro–Wilks for normality tests, asymmetry and
kurtosis. Data were analyzed with the ANOVA and
multiple comparisons by the Tukey test (p B 0.05).
The number of umbels per plant (17–25), capsule per
umbel (2–179) and per plant (554–1990), normal seed
per capsule (0–297), normal seeds per plant
(30,610–186,209) and sterile seeds per plant
(211,059–619,251) widely and significantly varied
among infructescences. Biomass of capsules per plant
umbel (3–795 g), biomass of sterile and normal seeds
per capsule (0.071–1.449 and 0–3.320 g), per umbel
(0.34–97.76 and 0.21–185.26 g) and per plant
(182–1052 and 334–2069 g) also varied widely. Seed
yield was statistically different between plants simul-
taneously growing and maturing at the same site.
Keywords Agave salmiana  Capsule 
Infructescence  Maguey  Reproduction by seed 
Umbel
Introduction
The Agave genus includes about 200 species; 75% of
them are distributed in Mexico (Garcı́a-Mendoza
2011; Reynoso-Santos et al. 2012). Agaves are long-
lived semelparous plants, which produce an
123

inflorescence only once, and at the end of its life cycle
the infructescence contains several thousands of
viable and sterile seeds (Peña-Valdivia et al. 2006;
Ramı́rez-Tobı́as et al. 2012). Therefore, agave plants
could spread sexually, by seeds. However, in situ
propagation is mainly asexual (vegetative shoots and
bulbils); which promotes initial seedling establish-
ment by dependence on the mother plant.
In Mexico, probably for centuries, Agave salmiana
plantations have been established with shoots of young
rhizomes (Mora-López et al. 2011). In the Sonora
Desert, from 1.2 million A. deserti seeds only one
develops a mature plant (Jordan and Nobel 1979). The
agave diversity is at risk due to its susceptibility to
pests and diseases of commercial clonal populations,
climatic change, and reductions in their geographic
distribution and number of wild populations. Multi-
plication via seeds increases genetic variability and
may also diminish damage resulting from pests and
diseases (Moreno 2003; Ramı́rez-Tobı́as et al. 2012;
Valenzuela-Zapata and Nabhan 2003).
In agaves seed production widely varies among
species, between 777 in A. angustifolia to 780 000 in
A. palmeri (Howell and Roth 1981; Molina-Freaner
and Eguiarte 2003). According to Reyes (2013), an
inflorescence of agave can produce more than 3000
seeds. Nobel (1998) estimated 65,000 seeds by
infructescence of A. deserti Engelmann, and Lorenzo
(2012) estimated 28,084 and 80,504 by infructescence
of A. salmiana subsp. crassispina (Trel. ex L. H. Bai-
ley) Gentry. The abundance of seeds per plant is also
been reported in A. americana L., A. angustifolia
Haw., A. celsii Hook. var. albicans (Jacobi) Gentry, A.
chrysoglossa I.M. Johnst., A. difformis A. Berger, A.
fourcroydes Lem., A. horrida Lem. ex Jacobi, A.
macroacantha Zucc., A. marmorata Roezl, A. mck-
elveyana Gentry, A. palmeri Engelm., A. tequilana
F.A.C. Weber, A. striata Zucc., A. subsimplex Trel., A.
vilmoriniana A. Berger and A. xylonacantha Salm-
Dyck (Arizaga and Ezcurra 2002; Arizaga et al.
2000a, b; Colunga-Garcı́aMarı́n and May-Pat 1997;
Colunga-Garcı́aMarı́n et al. 1996; Eguiarte et al. 2000;
Escobar-Guzmán et al. 2008; Howell and Roth 1981;
Lorenzo 2012; Molina-Freaner and Eguiarte 2003;
Nobel 1977; Rocha et al. 2005; Szarek and Holmesley
1996). But, in most of these studies seeds were not
directly accounted for, so these values may be
inaccurate. In this regard, according to information
in the literature, in 69.2% of the studies, the total
123
Genet Resour Crop Evol
number of seeds per plant was estimated from the
number of seeds counted in a small capsule sample; in
23.1% of them the calculation was partial, since the
counted seed number was from a small capsule
sample; and 7.7% of the studies did not specify how
the values were estimated.
The scarce number of studies concerning seed
production in A. salmiana is due to several reasons,
some of which are: this species, although widely
distributed, is endemic to Mexico, plants bloom only
once in their life and then die (monocarpic plants)
(Garcı́a 2017) and produce only one inflorescence per
plant, which is exposed when the plant is between 8
and 25 years old (Castro-Dı́az and Guerrero-Beltrán
2013); besides, the infructescence located at the apex
of the floral peduncle is at several meters above the
base of the plant (Garcı́a 2017; Granados 1993). Also,
fruits (capsules) are dehiscent (Garcı́a 2017) and
frequently not accessible because wild plants develop
in rugged topography sites (Lorenzo 2012); however,
overexploitation of different plant structures before
and during flowering cause the fruit absence (Garcı́a-
Mendoza 2011; Garcı́a-Moya et al. 2011). In this
regard, Lorenzo (2012) pointed that out of 100 plants
selected to evaluate the reproductive success of A.
salmiana, none produced infructescence because the
nearby villagers harvested all inflorescences.
The objective of this study was to quantify the yield
of capsules and seeds of A. salmiana Otto ex Salm-
Dyck plants simultaneously maturing and growing in
the same region. We hypothesized that, regardless of
the distribution of the capsules along the stalk, seed
yield per plant is similar between plants simultane-
ously maturing and growing in the same region.
Materials and methods
Collection site
Three plants of A. salmiana in reproductive stage, with
mature infructescences (Fig. 1) were found and har-
vested at San Luis Potosı́, SLP, Mexico (22°400 and
21°570 N, 100°440 and 101°110 W), at an altitude of
1870 m and BSokw (e)gw00 climate, equivalent to dry
steppe (BS), (o) arid, (k) temperate climate, (w) with
summer rains, (e) extreme climate with annual oscil-
lation of the mean monthly temperatures between 7
and 14 °C, (g) with Ganges-type temperature, and
Genet Resour Crop Evol
Infructescence
Stalk
Dead rosette
tissue
Valves
Fig. 1 Structures of an 8 m height plant of Agave salmiana Otto ex Salm-Dyck in fructification
(w00 ) with heat stroke, according to the Köppen
modified by Garcı́a (1988) classification (INEGI
2009).
Plant material
The mature infructescences, with unopened capsules,
were harvested between August and September 2014.
Each umbel was numbered in the acropetal direction
and separated from the infructescences. The capsules
of each umbel were allowed to dry for 4 weeks at room
temperature, in a dry, aerated and shaded place. They
were then stored in paper bags and kept in cardboard
boxes until evaluated (Fig. 1).
Evaluated variables
Capsules per umbel, normal and sterile seeds per
capsule and per umbel were counted. The diameter
and length of the capsules were measured with a
digital vernier caliper (Truper, CALDI-6MP; 14388).
The biomass (g) of capsules, valves per capsule and
per umbel, normal seeds per capsule and per umbel,
sterile seeds per capsule and per umbel were obtained
Umbels with capsules
capsules
Fruit
Sterile Normal
seeds seeds
on an analytical balance (accuracy of ± 0.0001 g;
ScientechÒ SA 120).
The number of sterile seeds was quantified in 11%
of the total capsules per umbel, distributed at the base,
center and apex of the infructescence. The percentage
corresponded to 59, 75 and 197 capsules per plant. The
biomass of the sterile seeds of each capsule was
obtained on an analytical balance (accuracy of ±
0.0001 g; ScientechÒ SA 120). The number of sterile
seeds per capsules, in which it was not counted, was
calculated by its biomass.
The seed moisture content was determined via
lyophilization, for 96 h, after freezing in liquid
nitrogen; six replicates per umbel, with 10 seeds each,
were evaluated. Seeds for this evaluation were
obtained from capsules of six umbels of the mid-
section of each infructescence.
Experimental design and statistical analysis
The study developed following a completely random-
ized design, with a plant as an experimental unit.
Frequency distribution of capsules number and
biomass, number and biomass of normal and sterile
123

seeds, and biomass of valves by umbel along the
infructescence stalk were tested for normality based
on the graphic residual analysis and the Shapiro–
Wilks test with the STATGRAPHICS software.
Variables were transformed when the normality
assumption was not met. The data were analyzed with
the ANOVA, Tukey’s multiple comparison tests and
Pearson’s correlation. Statistical analyses were per-
formed with the SAS statistical software (version 9).
Results and discussion
Number of umbels
Plants were randomly identified as plant 1, 2 and 3.
They had a total of 25, 17 and 19 umbels each along
the stalk.
Number of capsules
Plant 1, 2 and 3 had 1990, 590 and 554 total capsules,
respectively. The number of capsules per umbel in the
infructescence, in acropetal direction, was plotted to
know its distribution on the umbels along the
infructescence (Fig. 2). This showed a wide variability
(coefficient of variance: C.V. = 64.32, 45.86 and
57.32% in plant 1, 2 and 3 each) among umbels and
among plants. In plant 1 the frequency distribution of
capsules per umbel along the infructescence showed a
symmetrical trend around the mean. Thus, the first
umbel at the infructescence base and those on the apex
(Nos. 24 and 25) had the smallest amounts of capsules
Fig. 2 Number of capsules
per umbel, in acropetal
direction, in the
infructescence of three
Capsules per umbel (No.)
plants (A–C) (C.V. = 64.32,
45.86 and 57.32%) of Agave
salmiana Otto ex Salm-
Dyck
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Genet Resour Crop Evol
(six and 19). In contrast, the largest amounts of them
(172 and 179) were in some of the central umbels
(such as Nos. 12 and 14) (Fig. 2A).
Although basal and apical umbels (Nos. 1, 2, 16 and
17) in plant 2 infructescence had the lowest number of
capsules than in the central section, capsules distribu-
tion in umbels was not symmetric around the mean,
without the dominance of any umbel by capsule
content. Therefore, several umbels, with different
location, showed the highest similar amounts (49 and
55) of capsules (Fig. 2B).
The number of capsules per umbel in plant 3
distributed in the acropetal direction, was asymmetric
in the infructescence; the greater number of capsules
was concentrated in the umbels at the first two thirds,
from the base, of the infructescence. In contrast, the
three apical umbels (Nos. 17–19) had only 2% of the
total capsules of this plant (Fig. 2C).
These results showed differences in the number of
umbels and yield of capsules between the three plants,
which grow at the same site and matured in the same
period. In addition to the difference in capsule yield,
contrasts in capsule distribution on the umbels and on
infructescence stood out. Regardless of the number of
umbels in the plants, those on the three plant apex
showed lower capsule production and only those at the
base of two infructescences also showed low capsule
amount; but their distribution along the infructescence
was uneven. This tendency was partially similar to that
registered in infructescences of A. salmiana in San
José Alchichica, Puebla, where the umbels with the
highest capsule production were found at the first half
of the infructescence (Lorenzo 2012).
Base Apex Base Apex Base Apex
200
(A) (B) (C)
150
100
50
0
0 5 10 15 20 25 0 5 10 15 20 25 0 5 10 15 20 25
Number of umbel on the stalk
Genet Resour Crop Evol
The total number of capsules was significantly
different (p B 0.05) between plants. Plants 2 and 3
showed 70 and 72% fewer capsules compared to plant
1. Lorenzo (2012) obtained similar differences when
comparing two infructescences of A. salmiana, with
the production of 75 and 277 capsules. Those varia-
tions between infructescences are due to differences in
the number of pollinated flowers in each umbel and in
the complete inflorescence, but also to the abortion of
immature fruits due to the number of available
resources, flower and fruit herbivory, among other
factors (Arizaga and Ezcurra 1995; Ehrlén 1991;
Stephenson 1981).
The frequency in the distribution of the number of
capsules per umbel showed a significant drop in their
number after the first third, with an acropetal direction,
in the three infrutescences (Fig. 2). These trends seem
to be a multifactorial pattern. Among factors are those
intrinsic to plants, such as the synchrony in the
resources allocation from the stalk to the formation of
reproductive structures and the second order (or
branches) for a similar photo-assimilate remobiliza-
tion. Others are environmental factors that affect both
the rate of inflorescence exposure and flower and fruit
abscission. Another is the pollination that, at the same
time, depends on multiple factors (Trejo-Salazar et al.
2015). It should be noted that the inflorescence in
agaves, like those of other species, is a physiologically
fragile structure, but exposed to environmental effects
during flowering period for relatively long periods of
15 days or more (Rocha et al. 2005).
Diameter and length of the capsules
The mean diameter of capsules in plant 1 was
22.50 mm (n = 1988) and was significantly higher
(p B 0.005) than in plant 2 (21.25 mm with n = 589)
and plant 3 (20.15 mm with n = 554); the difference
was between 5.56 and 10.44% between plant 1 and the
other two. This capsule characteristic had more
variability in plant 1 (C.V. = 10.18%) than in the
others (C.V. = 7.77 and 6.18%). The capsule diameter
at the present study was in the interval (20–24 mm)
obtained for A. salmiana plants at the Valley of
Mexico (Rzedowski and Rzedowski 2005) and the
Tehuacán-Cuicatlán Valley, Mexico (20–30 mm)
(Garcı́a-Mendoza 2011).
Regardless of the mean diameter of the capsules,
these structures appeared heterogeneous, and small,
medium and large capsules were identifiable. So the
mean capsule diameter per umbel of each plant was
plotted to know how it changes along the infructes-
cence. None of the plants showed any direct trend in
the diameter of the capsule with respect to the position
(or number) of the umbel in the infructescence; in most
umbels there was some significant difference in the
mean capsule diameter. The C.V. showed that the
capsule diameter variability on the umbels of plant 1
(from 5.70% in the first umbel to 13.86% in No. 20),
plant 2 (from 3.86% in umbel No. 5 to 11.05% in No.
10) and plant 3 (from 5.69% in umbel 1 and 16.66% in
No. 19) was similarly heterogeneous (Fig. 3). Col-
unga-Garcı́aMarı́n et al. (1996) and Colunga-Garcı́a-
Marı́n and May-Pat (1997) documented little
variation, based on the standard deviation (S.D.), on
the mean capsule diameter between A. angustifolia
and A. fourcroydes.
Some significant differences (p B 0.05) on the
mean diameter of the capsules per umbel existed on
each plant. The smaller diameter capsules (20.75 and
20.71 mm) in plant 1 were located at the central umbels
(Nos. 11 and 12), the largest ones (22.22 and
22.25 mm) in the umbels close to the infructescence
apex (Nos. 17 and 19) (Fig. 3A). In plant 2, mean
capsule diameter was also significantly different
among few umbels; in fact, umbel No. 16, at the apex
of the infructescence, had capsules with a significantly
smaller mean diameter (19.64 mm) than both at the
base of infructescence (21.74 and 21.51 mm)
(Fig. 3B). In plant 3 there were umbels with low mean
diameter capsules (16–19 mm) near the base (umbel
No. 3) and at the apex (umbel Nos. 15 and 19) of the
infructescence. In this plant, the largest mean diameter
(21.49 mm) was significant (p B 0.05) only on the
middle umbel (No. 8) of the infructescence (Fig. 3C).
The mean length of the capsules was significantly
different (p B 0.001) between plants; although, this
difference amounted only 1.8%. In plant 1 the capsule
mean length was lower (54.30 mm, n = 1988) than in
plant 2 (54.92 mm, n = 589). The mean length of plant
3 capsules was the lowest (45.68 mm; n = 554) and
had 16 and 17% less length than in the other plants.
The mean length of the capsules in this study was
(46–55 mm) similar to that reported for A. salmiana
growing in North America and the ‘‘Valle de México’’
(55–70 mm) (Gentry 1972; Rzedowski and Rze-
dowski 2005) and at the Tehuacán-Cuicatlán Valley,
Mexico (60–70 mm) (Garcı́a-Mendoza 2011).
123
 Genet Resour Crop Evol

Base Apex Base Apex Base Apex

24
(A) (B) (C)

Capsule diameter (mm)

* * **
22 *
**
* *
20 *
*

18 *

16

(D) (E) (F)

Capsule lenght (mm)

60 * ** *
* * * * *
* *
** *
50 * * ** *
*
* *
40 *

30
0 5 10 15 20 25 0 5 10 15 20 25 0 5 10 15 20 25
Number of umbel on the stalk

Fig. 3 Mean diameter of the capsules (? DE), in acropetal
direction, in the umbels of the infructescence of three plants (A–
C) (CV of 5.70–13.86, 3.86–11.05 and 5.69–16.66%) and length
of capsules (? DE) on the umbels of the infructescence, in
acropetal direction, of three plants (C–E) (CV of 6.03–21.67,
6.57–10.92 and 7.21–13.22%) of Agave salmiana Otto ex Salm-
Dyck. Asterisks on bars indicate statistical differences (p
B 0.05) between that mean and at least one other (plant 1:
white bars, plant 2: light gray bars and plant 3: dark bars)

The mean length of the capsules was plotted by Biomass of capsules and valves
umbel to visualize their distribution along the
infructescence. In plant 1 a certain gradient was The total capsule biomass by infructescence was
observed in said length. The longest capsules statistically different between plants (p B 0.05); plant
(56–57 mm) corresponded to the umbels in the first 1 (8 750 g) had 3.6 and 5.6 times greater biomass than
basal third of infructescence and the shortest ones plants 2 (2 467 g) and 3 (1 567 g). In contrast, the
(46–47 mm) in the apical region; but, these differ- mean capsule biomass in plant 1 (4.40 g) was
ences were significant (p B 0.05) only between some statistically similar than plant 2 (4.18 g) (p = 0.151),
umbels (Fig. 3D). This trend was similar in plants 2 and both were almost twice (p B 0.05) than plant 3
and 3, which the capsule mean lengths on umbels (2.66 g). The C.V. (34.51, 26.3 and 43.3% in plants 1,
(Nos. 2–5) at the base of the infructescence were 2 and 3) indicate that this characteristic was remark-
around 56 and 49 mm and those near to the apex were ably variable within the plants.
55–45 mm or less (Fig. 3E, F). The capsule length was The mean biomass of the capsules by umbel varied
heterogeneous between the umbels of the three plants, between plants and showed no relation to the location
since their C.V. was 6.03–21.67, 6.57–10.92, and of umbel on infructescence. Also, the C.V. of capsule
7.21–13.22% in the umbels of plant 1, 2 and 3 (Fig. 3). biomass by umbel was one of the most variable
These results indicate that the maximum longitudinal characteristics of the infructescence. In plant 1 the
growth of the capsules in the umbels is remarkably C.V. of this characteristic ranged from 26.86 to
heterogeneous with a similar growth pattern between 75.30% (in umbel Nos. 1 and 24), in plant 2 from
plants. 24.17 to 34.81% (in umbel Nos. 9 and 17) and plant 3
from 25.98 to 57.14% (in umbel Nos. 1 and 16).

123
Genet Resour Crop Evol
Irrespective of this variability, the differences between
some of the umbels were significant (p B 0.05), but
without any direct trend (Fig. 4A–C). Likewise cap-
sule length, the mean capsule biomass, at the end of
infructescence growth, was remarkably heterogeneous
between umbels in each plant, with a different pattern
among plants; nevertheless, it seems to be associated
to the number of capsules per umbel.
It is unknown whether the growth and development
of capsules along the stalk shows any tendency regard
the umbel position which they are on; this information
may indirectly indicate patterns of nutrient mapping
throughout the stalk. The distribution of the total
biomass of the capsules by umbel along the stalk,
described a Chi-square tendency (p B 0.05), non-
Gaussian, in plant 1. Although this plant had umbels
intercalated with low biomass, out of trend, those with
the highest total capsule biomass were those located in
the middle section of the infructescence (Fig. 5A).
The distribution of the total biomass of capsules per
umbel in the infructescence of plant 2 showed a
partially similar tendency to that described on plant 1
because the umbels at the base and the apex capsules
biomass were lower than in most of the central ones.
But, the mean capsule biomass distribution in the
umbels of this plant was multimodal, with more than
two maxima (Fig. 5B). In contrast to these two plants,
the total biomass distribution of the capsules per
umbel in the infructescence of plant 3 showed an
asymmetric and truncated tendency. This distribution
showed that in the first two-thirds of the infructes-
cence, in the acropetal direction, the capsule biomass
production was relatively constant, except for umbel
Fig. 4 Mean biomass of the Base
capsules (? D.E.) on
umbels of the 7 (A)
infructescences, in acropetal
direction, of three plants (A– 6
C) of Agave salmiana Otto
5 **
Capsule (g)
ex Salm-Dyck. The asterisks
on bars indicate statistical
4 *
differences (p B 0.05)
between that mean and at 3 *
least one other (plant 1:
white bars, plant 2: light 2 *
gray bars and plant 3: dark
bars) 1
0
0 5
No. 5; the biomass of capsules in the last apical third
was significantly lower than in the rest of the umbels
(Fig. 5C).
The total biomass of the valves in plant 1 (5530 g)
was 3.23 and 5.22 times greater than in plants 2
(1711 g) and 3 (1059 g) (Fig. 6). The valves biomass
distribution on the umbels was remarkably similar to
that of the capsules. This distribution on the umbels,
along the stalk, in the plant 1 was of the Chi square
type (p B 0.05), and multimodal with more than two
maximum values in plant 2 and asymmetric and
truncated in plant 3 (Fig. 5D–F).
Differences in the total biomass of capsules and
valves are due to the fact that in the second case seeds
were extracted (Fig. 5). These differences allow us to
anticipate that the seeds represent, on average, about a
third of the total biomass of the capsules, indepen-
dently of the plant and umbel to which they belong.
Number of seeds
The three plants showed high sterile seed content. The
mean number of sterile seeds per capsule was 311, 369
and 364 in plant 1, 2 and 3 each; and in plant 1 it was
significantly lower (p B 0.05) than in the other two.
The number of sterile seeds in the three plants was
higher than those reported for A. angustifolia
(119–168 and 140–195), A. fourcroydes (220–281),
A. macroacantha (98.68, 78.65 and 102.97), and A.
salmiana (187–237) (Arizaga et al. 2000b; Arizaga
and Ezcurra 2002; Colunga-Garcı́aMarı́n et al. 1996;
Colunga-Garcı́aMarı́n and May-Pat 1997; Lorenzo
2012). Probably due to the total number of seeds in
Apex Base Apex Base Apex
(B) (C)
*
*
* *
* ** * *
*
*
* * * * ***
10 15 20 25 0 5 10 15 20 25 0 5 10 15 20 25
Number of umbel on the stalk
123
 Genet Resour Crop Evol

Fig. 5 Total biomass of Base Ápice Base Ápice Base Ápice

capsules (A–C) (CV 60.63,
64.32 and 65.27%) and 800 (A) (B) (C) 800
valves (D–F) (CV 46.17,
45.86 and 62.04%), in

Capsules (g)
acropetal direction, by 600 600

Capsules (g)
umbel of infructescence of
three plants of Agave
salmiana Otto ex Salm- 400 400
Dyck (plant 1: white bars,
plant 2: light gray bars and
plant 3: dark bars) 200 200

0 0
800 (D) (E) (F) 800

600 600

Valves (g)
Valves (g)

400 400

200 200

0 0
0 5 10 15 20 25 0 5 10 15 20 25 0 5 10 15 20 25
Number of umbel on the stalk

maguey plants, most studies only estimate the number
of sterile seeds, so, those values could be imprecise.
Counting sterile seeds is difficult because, in
addition to their abundance in the capsules, they are
in packaged rows of several tens and mixed with
normal seeds (Fig. 7). For this reason, sterile seeds of
all capsules of three umbels (at the base, center, and
apex) per infructescence were counted. In plant 1
those umbels were Nos. 1, 12 and 25. The umbels that
were evaluated in the other plants were different,
except for the basal, because the number of umbels
was different between plants (Fig. 2). The umbels
evaluated in plant 2 were Nos. 1, 9 and 17 and in plant
3, in addition to the first one, Nos. 10 and 19 were
evaluated. Thus, the total capsules used for a direct
count of sterile seeds were 197, 59 and 74 of plant 1, 2
and 3. The total number of seeds counted directly in
the plants was 57,416, 22,353 and 27,673, respec-
tively. With this information and the seed biomass, the
number of sterile seeds was calculated in all umbels of
each plant.
The counts and calculated values of the numbers of
sterile seeds in the rest of the umbels indicated that
plant 1, 2 and 3 could contain around 619,251, 217,506
and 211,059 of these seeds. This indicated that plants 2
and 3 would have a third than plant 1. This quantity of
sterile seeds in the infructescences was between 100
and 300 times greater than that estimated for A.
macroacantha (1887), and seven to 34 times more
than the amount calculated for A. salmiana
(18,330–29,126) (Arizaga and Ezcurra 2002; Lorenzo
2012).
Because the umbels on the central zone of each
infructescence contained a significantly larger
amounts of capsules than those on the base and apex,
the larger number of sterile seeds showed a Chi-square
distribution in plant 1 (p B 0.05) and a Gaussian bias
to the right on plant 2 (p B 0.05) (Fig. 8A–B). In
contrast, the larger amounts of sterile seeds in plant 3
were in the capsules on umbels of the base and those
up to the first two thirds of infructescence compared to
the apex. Seed frequency distribution by umbel
showed two maxima and their frequency showed
asymmetric and truncated tendency (Fig. 8C).
The number of normal seeds of the three plants was
obtained by direct counting. In plant 1 was 186,209;
which was more than six and five times higher than in
plant 2 and 3 (30,610 and 35,882 each). The normal

123
Genet Resour Crop Evol

(A) seed distribution on umbels along the infructescence

was remarkably similar to that described for sterile
5 530 g Valves
seeds, with a Chi square type in plants 1 and 3
2 069 g Normal seeds
1 052 g Sterile seeds (p B 0.05), and of a Gaussian type in plant 2
(p B 0.05) (Fig. 8D–F). In plants 1 and 2 most of
8 651 g Biomass of capsules
per plant
the central umbels had larger numbers of capsules, and
therefore normal seeds tended to concentrate in some
of these umbels, similarly to sterile seeds. In the plant
1 case, the first four umbels on the base contained
(B)
between 1 743 and 8 304 normal seeds and the six most
1 711 g Valves apical umbels contained between 274 and 1 659 of
380 g Normal seeds these seeds; in contrast, some of the central umbels
362 g Sterile seeds
(12–17) had between 10,700 and 16,662 normal seeds
2 453 g Biomass of capsules (Fig. 7D). A similar trend was observed in infructes-
per plant
cence of plant 2, with about 500 normal seeds on the
basal and apical umbels and more than 3000 and 1500
normal seeds between the third and fifteenth umbel
(C) (Fig. 7E). In contrast, the first 11 umbels of plant 3
1 059 g Valves
accumulated 30 563 normal seeds, which represented
334 g Normal seeds 85.18% of the whole plant. These results showed that
182 g Sterile seeds the number of sterile seeds respect the normal ones
1 575 g Biomass of capsules was greater in the three plants, but normal seeds of
per plant plant 1 represented 31.28% of the sterile seeds, and in
plant 2 and 3 that proportion was only 13.88 and
17.00%.
To confirm whether these proportions of normal
Fig. 6 Proportion of valves, sterile and normal seeds by (counted) and sterile seeds (for which a part was
infructescence of three plants (A–C) of Agave salmiana Otto
ex Salm-Dyck
calculated) were attached to the actual values, the
percentages were calculated only with data from the
umbels (Nos. 1, 12 and 25 in plant 1, Nos. 1, 9 and 17
in plant 2, Nos. 1, 10 and 19 in plant 3 in the
acropetally direction) where both types of seeds were
directly counted. Thus, the number of normal seeds
represented 31.89% of the sterile seeds in the plant 1,
14.39 and 15.80% in the plants 2 and 3. The values
were similar to those calculated for the whole
infructescence, and are evidence that the calculated
number of sterile seeds for the three plants was correct.
In contrast to these seed ratio, estimates of sterile seeds
in A. macroacantha and other A. salmiana plants
accounted for only half of the normal seeds (Arizaga
and Ezcurra 2002; Lorenzo 2012).

Seed biomass

The total biomass of the sterile seeds in plant 1 was

Fig. 7 The arrangement of normal seeds (black) and sterile about three times greater (p B 0.05) than in plant 2
(light color) in a capsule of Agave salmiana Otto ex Salm-Dyck
and six times that of plant 3. At the same time, the
biomass of the normal seeds in plant 1 represented

123

Base Apex
50000
(A)
40000
30000
20000
10000
Seeds (Num.)
0
(D)
15000
10000
5000
0
0 5 10 15 20
Fig. 8 Total number of sterile (A–C) seeds (CV 67.92, 66.84
and 62.71%) and normal (D–F) seeds (CV 48.80, 48.89 and
78.48%) by umbel, in the acropetal direction in the
about 5.5 times that of plant 2 and more than 6.2 times
from plant 3 (Fig. 6).
Despite differences in biomass of capsules by
umbel or infructescence in all plants, the proportion of
sterile and normal seeds was similar among plants.
The biomass of sterile seeds represented 12, 15 and
12% of the capsules in plant 1, 2 and 3; the relative
proportion of normal seeds was 24, 16 and 21%, and
therefore, valves biomass also had relatively similar
proportions (64, 70 and 67%) among them (Fig. 6).
In general, sterile and normal seed biomass in
plants 1 and 2 was low in basal and apical umbels.
Although the biomass distribution of both seed types,
on each umbel of the three plants was heterogeneous,
central umbels of the infructescences accumulated
larger amounts of both types of seeds. In plant 1, this
trend was evident; because 26% of the total biomass of
sterile seeds was accumulated in three of the 25
umbels (Nos. 12, 14 and 15) and, even more, four
umbels (Nos. 12–15) concentrated 34% of the total
123
Genet Resour Crop Evol
Base Apex Base Apex
(B) (C)
(E) (F)
25 0 5 10 15 20 25 0 5 10 15 20 25
Number of umbel on the stalk
infructescences of three plants of Agave salmiana Otto ex
Salm-Dyck (plant 1: white bars, plant 2: light gray bars and plant
3: dark bars)
biomass of normal seeds in infructescence of plant 1
(Fig. 9A, D). In plant 2 between seven and nine central
umbels (Nos. 3, 5–7, 9, 11–15) accumulated the
largest proportions of both seed types (Fig. 9B, E). In
contrast to plants 1 and 2, plant 3 showed the highest
biomass of both seed types (94.35% sterile and
92.21% normal seeds) on the base and central umbels
(Nos. 1–13) of the inflorescence. Umbel No. 12 of this
plant showed an atypical high normal seed biomass,
with up to 300 times more seed biomass respect to
umbels Nos. 5 and 19 (Fig. 9C, F). These results
indicate that the biomass distribution of both groups of
seeds in the umbels, throughout the infructescence,
was partially different between plants, but similar
within each plant.
Seed moisture
The moisture content in the normal seeds of three
plants was statistically similar (p = 0.1308;
Genet Resour Crop Evol
Base
100
(A)
80
60
40
Biomass of seed (g)
20
0
(D)
150
100
50
0
0 5 10
Fig. 9 Biomass of sterile (A–C) (CV 66.14, 68.20 and 62.82%)
and normal seeds (D–F) (CV 47.88, 48.34 and 79.23%) by
umbel, in the acropetal direction in infructescences of three
7.18 ± 0.55, 6.54 ± 0.33 and 6.42 ± 5.94%) among
them. In contrast, Vazquez et al. (2011) determined
that the seeds of the Blanco, Chino and Liso variants of
A. salmiana, harvested in San Luis Potosı́, had
different moisture content, between 2.8 and 11.6%.
However, seed moisture depends on the environment
in which they are at the end and after maturation. The
low moisture content in agave seeds, as in other
species, may favor the maintenance of their viability
(Doria 2010) at in situ seed bank.
To estimate the number of sterile seeds, the
assessment of their biomass seems appropriate since
a positive due to the correlation between the number
and weight of these seeds in plants 1, 2 and 3
(r = 0.9376, 0.9480 and 0.997). Another option is to
use the fruit weight because this variable was also
positively correlated with the number of sterile seeds
(r = 0.7501 for plant 1, r = 0.8364 for plant 2, and
r 0.975 for plant 3; p B 0.05). In this study moisture
content of seeds did not affect the difference of the
total seed biomass between the plants.
Differences in the umbels amount and distribution
capsules per plant and umbel, and viable seeds per
capsule and per plant documented in the literature and
Apex Base Apex Base Apex
(B) (C)
(E) (F)
15 20 25 0 5 10 15 20 25 0 5 10 15 20 25
Number of umbel on the stalk
plants of Agave salmiana Otto ex Salm-Dyck (plant 1: white
bars, plant 2: light gray bars and plant 3: dark bars)
in the present study may be due to plant internal
factors, such as their genotype, and external, as the
environment plants are growing. In addition, in
relation to seeds, one of the factors responsible for
the differences between genotypes, both in the liter-
ature and the results here presented must partially
responded to the natural pollinators such as bats
(Corynorhinus townsendii), bees (Apis mellifera),
hummingbirds (Colibri sp.), wasps (Hymenoptera),
butterflies (Lepidoptera) and ants (Solenopsis saevis-
sima) (Howell and Roth 1981; Granados 1993).
In situ sexual multiplication of agaves depends on
factors such as pollinators, seed dispersion, seed and
seedling predators, conditions for germination and
seedling survival, which can be restrictive in natural
environments within their distribution (Garcı́a-Men-
doza 2007; Mora-López et al. 2011). In spite of limited
natural sexual multiplication, persistent maguey cap-
sules frequently contain abundant seeds (Moreno
2003; Garcı́a-Mendoza 2007). The Agave seed dis-
persion is favored by its inflorescence elevated
position in the landscape (up to several meters above
the ground; Fig. 1) and its shape (flattened) with a
short wing in its convex portion (Garcı́a-Mendoza
123

2011). However, some species, such as A. schottii
Engelm., only disperse 15% of their seeds (Trame
et al. 1995).
Conclusions
The number of seeds in A. salmiana plants may be
independent of the growth site and maturation period
of the infructescence. Regardless of the number of
umbels and capsules per umbel, the distribution of the
capsules along the infructescence was asymmetric.
The greatest capsule quantity and consequently of
normal and sterile seeds can be located in umbels at
the base and centers of the infructescence. The high
number of sterile seeds indicates that many maguey
seeds do not develop completely, and remain sterile;
agave seeds viability is a complex process that
requires further studies, given that the assigned plant
resources to these structures are abundant.
Acknowledgements The authors would like to thank M.Sc.
Victor Baruch Arroyo Peña for the critical review and
improvement for this version of the document.
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