TAXON 27(5/6): 463-470.
NOVEMBER 1978
STRUCTUREOF STARCHGRAINSAND THE CLASSIFICATIONOF
VASCULARPLANT FAMILIES
A. Th. Czaja*
Definition of some vascular plant families is unsatisfactory and, consequently, it is impossible to classify them adequately. The lack of an analytical key to
the families based on easily observed, widespread characters sometimes results
in surprising misinterpretations.
A striking example in this regard is the Casuarinaceae. Engler (1897) considered them to be true Angiospermae (Dicotyledoneae, Order 1. Verticillatae).
Melchior (Engler's Syllabus der Pflanzenfamilien, vol. 2. 1964) and Wettstein
(1935) concur with Engler, but Hutchinson (1973) placed the Casuarinales
(Order 18) between the Juglandales and Urticales. According to Takhtajan
(1969, 1973) the Casuarinaceae probably arose from the Hamamelidales. Gaussen (1940) derived the Casuarinaceae from the Articulatae. Hegnauer (1964)
recalls that authors have noted similarities between Casuarina, Equisetum, Ephedra, and the Coniferae, but states the chemistry of the family is not well
enough known to determine its position in the system.
The Nymphaeaceae is another difficult family to place in the system. In
Engler's Syllabus, Order 16 Ranunculales includes the suborders Ranunculineae
and Nymphaeineae. The latter contains families Nymphaeaceae, and Ceratophyllaceae. The first of these is divided into the subfamilies Cabomboideae,
Nymphaeoideae and Nelumbonoideae. Wettstein places the Nymphaeaceae in
Order 18, Polycarpicae, following Family 20, Berberidaceae. The Nymphaeaceae contains the same subfamilies as in Engler's Syllabus. The following
family is Ceratophyllaceae, which is considered closely related to the Nymphaeaceae. Hegnauer (1969) states that the evidence is not sufficient to form an
opinion with respect to the relationships of all the genera included in the
Nymphaeaceae. Hutchinson puts the family in the Ranales and it includes
Nelumbo, Euryale, Nuphar and Barclaya. He treats Cabombaceae and Ceratophyllaceae as separate families. According to Kubitzki (1972), the Nymphaeaceae, s. str., do not belong in the Polycarpicae.
A third example is the Cyclanthaceae. Wettstein placed them in the Order
Synanthae, with the Palmae and Araceae but the plants of the three families are
very different. The Palmae are mostly derived, the Cyclanthaceae include at
least two different plant types, and the Araceae, mostly herbaceous, exhibit
distinctive features. In Engler's Syllabus, the Order Synanthae (Cyclanthales)
comprises the family Cyclanthaceae with subfamilies Carludovicoideae and
Cyclanthoideae. These are herbs of palm-like habit and deeply bilobed leaves.
They are very old, reduced and derived relicts. Hegnauer considers the Palmae,
Cyclanthaceae and Pandanaceae more closely related to one another than to
the herbaceous Araceae, Typhaceae and Sparganiaceae. According to Takhtajan, the Arales, Arecales and Cyclanthales were derived from the immediate
ancestors of the Liliales.
These three examples, as well as numerous others that could be cited, show
the lack of one or more constant characters which can be used to group
families into higher categories. The author believes the structure of starch
grains offers such characters. For more than thirty years he has conducted
microscopic identification of powdered plant materials, including foodstuffs,
flours, drugs and fibers. Microscopic examination of such substances in polar* Botanisches
Institut, RWTH Aachen. 51 Aachen, BRD.
NOVEMBER
1978
463
ized light is more informativethan in ordinarylight because the double refracting plant materialsreveal the submicroscopicstructure of their particlesunder
polarized light. Starch grains are particularlysignificant since they are ubiquitous in plants. The extraordinaryvariability of starch grains has been demonstrated by Nageli (1858), A. Meyer (1895), E. Reichert (1913) and many others. In his "Mikroskopieder Starkekbrner"(1969), the author has systematically examined starchgrainsto determinetheir submicroscopicstructureand their
distribution in the variousplant families. A number of basic construction types
of starch grains are distinguishableon sight; in addition, further differences
appearwhen the grains are observed carefully as they swell in a water preparation under a cover-glassover a pilot-light of a gas burner.
(1) The basic construction types of starch grainsare given in the chart below.
Because the terms applied to them are descriptive,arbitraryabbreviations
are given; the German equivalents are the terms used in my forthcoming
book on this subject.
Basic Construction Types
of Starch Grains
Grund- Bau- Typen
(Original Designation in German)
1 Envelope Starch Grains, round
(Envel.-St.-Gr., round)
Hiillen - Starke-Kbrner, rund
(Hii-St-K, rund)
2 Envelope Starch Grains, oval
(Envel.-St-Gr., oval)
Hiillen-Starke-Korner,oval
(Hii-St-K, oval)
3 Streched-Envelope Starch Grains, at first
round
(Str. Envel.St-Gr. first round)
Gestreckte Hi-St-K, anfangs rund
4 Highly Compound Starch Grains
(Highly Comp.-St!Gr.)
Hoch-zusammengesetzte St-K.
(Hoch zus. ges. St-K.)
5 Compound Envelope Starch Grains
(Comp.St-Gr.)
Zusammengesetzte St-K.
(Zus.ges. St-K.)
6 Cistaceae Envelope Starch Grains
(Cist.Envel.St-Gr)
Cistaceae-Hii-St-K.
(Cist.Hii-St-K.)
7 Papilionaceae Envelope Starch Grains
(Pap. Envel. St-Gr.)
Papilionaceae Hii-St-K.
(Pap.HiuSt-K.)
8 Papilionaceae (Mucuna) Envelope
Layer Starch Grains
(Pap.-Mucuna) Envel.-Lay.St-Gr)
Papilionaceae (Mucuna) Hii-St-K.
9 Shell Starch Grains (Triticeae)
formed in 2 steps, Endosperm
(Trit. Shell-St-Gr.)
Schalen-Starke-Kbrner(Triticeae)
(geb. in 2 Schritten, Endosperm)
(Trit.Schal.St-K.Endosp.)
(Gestr. anfangs runde Hii-St-K)
(Pap. (Mucuna) Hii-Lag.St-K.)
10 Pteridophyte Starch Grains
(Pter-St-Gr)
Pteridophyten-St-K.
(Pter.-St-K.)
11 Envelope Layer Starch Grains
(Envel.-Lay-St-Gr.)
Hiillen-Lagen-Starke-Korner
(Hu-Lag-St-K.)
464
TAXON VOLUME 27
12 Nearly Layer Starch Grains
13 15 (Nearly-Lay.St-Gr.)
Fast-Lagen-Starke-Korner
(Fast-Lag.St-K)
14 Layer Starch Grains
(Lay.-St-Gr.)
Lagen-Starke-Korner
(Layer-St-K.)
Starch Grains in the laticifers of
various and peculiar shapes (rods,
bones, dumbbells, sphaeroids) only
in some Euphorbiaceae
Starkekorner in Milchsaftschlauchen
in verschiedener und besonderer
Gestalt (Stabe, Knochen, Hanteln, Kugeln)
nur in einigen Euphorbiaceen
The numbers correspond to those in plate, I and II (abbreviations in parentheses).
The authenticity of these basic-construction-typesand their connections with
the families, orders or other systematic units, may probably be assuredby the
findings of M. Ludtke (1976). After the acetylation of potato, maize and rice
starch there is a residue with a complex of carbohydrateand protein. Hydrolysis with 2-n HCl produces glucose and maltose; a hydrolysis with 6-n HCl
produces the following amino-acids:asparaginicacid, threonine, serine, glutamic acid, glycine, alanine, ornithine, lysine. In addition arginine, leucine and
histidine were found in rice starch. The starch granulesare therefore described
as tissues, because these are not pure carbohydrates.
?oQf
0
?
J0D'+
O1 c
i
3
2
Sw
5CQP
0@0
5o
(
9?00
0
8
Plate I.
1. Envelope starch grains, round.
2. Envelope starch grains, oval (Quercus seeds, et al.).
3. Envelope starch grains, round, later stretched (Brominae).
4. High compound starch grains (Schizaeaceae, Festuceae, real Monocot., Centrospermae).
5. Compound starch grains (many lignif. plants, et al.).
6. Cistaceae: Envelope starch grains.
7. Papilionaceae: Envelope starch grains.
8. Envelope-layer starch grains (Mucuna seeds, Papilion.).
9. Shell starch grains (Triticeae).
NOVEMBER
1978
465
0
. .iII
+RF~~
~~IS~
11
10
14
13
+by+R
12o
+
$
15
Plate II
10. Pteridophytic starch grains.
11. Envelope-layer starch grains (Angiopteris, Marattiaceae, many Monocot. and Dicot.).
12. Nearly-layer starch grains (Orchidac. Phajus).
13. Nearly-layer starch grains (Curcuma, Zingib.).
14. Layer starch grains (Canna sp.).
15. Nearly-layer starch grains (Marattiales, Christensenia, Musa sp.).
Symbols:
(2)
o = ordinary light
+ = crossed Nicols.
+R = + and Red. I.O.
sw = swollen by heat.
The basic construction types of starch grains as they occur in divisions,
classes, orders, and families are significant.
PTERIDOPHYTA. The sequence of classes, orders and families is different from that in
Engler's Syllabus because of the many exceptions in some of the groups recognized there.
(a) Vegetative organs of all groups examined show starch grains of the Pteridophytic-B-CType. Following families examined: Psilotaceae, Lycopodiaceae, Equisetaceae, Isoetaceae, Salviniaceae (Salvinia, Azolla), Pilulariaceae, Marsileaceae (Marsilea, Regnellidium),
Ophioglossaceae (Ophioglossum, Botrychium), Osmundaceae, Schizaeaceae, Dicksoniaceae, Cyatheaceae, Polypodiaceae (s. lat.).
(b) Spores and microspores accumulate fatty oil and proteins. The megaspores of Isoetaceae, Salviniaceae, Pilulariaceae, and Marsileaceae contain starch grains in addition to
protein and fatty oil.
(c) Exceptions: Starch grains are quite absent in the Selaginellaceae; in some species trehalose has been detected as well as glucose and fructose but no sucrose.
In Equisetum and Osmunda, the spores are green, contain chloroplasts, but no
starch, are short-lived, must immediately germinate or die.
Vegetative organs of Danaea and some Marattia accumulate starch grains in the
parenchyma cells of the same Pteridophyte Type as the other pteridophyte families.
However, at the end of the recent Pteridophyta, within the Marattiales, other types of
starch grains occur; these are surely very old, for the Marattiales are undoubtedly relics
of the extinct fern vegetation of the Middle Carboniferous (Smith, G. M. 1955). Some
Marattia species (M. erecta, M. excavata, M. fraxinea, M. kaulfussii) have Envelope Layer
466
TAXON VOLUME 27
Type of starch grains. Christensenia aesculifolia and C. cumingiana have Nearly Layer
Type of starch grains and Angiopteris evecta also has Envelope Layer starch grains.
MONOCOTYLEDONES. The basic starch grain types among the monocots can be summarized in two columns, using the orders and families recognized in Engler's Syllabus II (1964).
Pteridophyte Type
Helobiae (Alismatales): Alismaceae
(Alisma, Limnophyton, Echinodorus,
Sagittaria); Butomaceae (Butomus,
Limnocharis)
Scheuchzeriaceae (Scheuchzeria)
Aponogetonaceae (Aponogeton)
Juncaginaceae (Triglochin, Lilaea)
Potamogetomaceae (Potamogeton)
Najadaceae (Najas)
Juncales: Juncaceae (Juncus, Prionium),
Lemnaceae (Lemna, Wolffia)
Mayaceae (Mayaca)
Restionaceae (Restio, Chondropetalum)
Pandanales: Pandanaceae (Pandanus), Sparganiaceae (Sparganium), Typhaceae (Typha)
Cyperales: Cyperaceae (Carex, Cladium,
Cyperus, Eleocharis)
Other Types
Hydrocharitaceae (Hydrocharis,
Vallisneria, Elodea, Stratiotes)
Liliiflorae: Liliaceaceae, Agavaceae,
Ilaemodoraceae, Cyanastraceae, Amaryllidaceae, Hypoxidaceae, Velloziaceae,
Taccaceae, Dioscoreaceae, Pontederiaceae,
Iridaceae, Burmanniaceae, Philydraceae
Bromeliales: Bromeliaceae
Commelinales: Commelinaceae,
Xyridaceae, Rapateaceae, Eriocaulaceae
Centrolepidaceae, Flagellariaceae
Graminales
Palmales
Cyclanthales
Arales
Scitamineae: Musaceae, Zingiberaceae,
Cannaceae, Marantaceae
From the standpoint of starch grains or carbohydrate substitutes, the Monocotyledones
families fall into three groups:
Primary Monocotyledones. Seeds and vegetative organs accumulate starch only in the
form of Pteridophyte Type of grains. This group, which is amylophilic, includes the
following families which have been examined: Alismaceae, Butomaceae, Juncaginaceae, Aponogetonaceae, Potamogetonaceae, Scheuchzeriaceae, Najadaceae, Restionaceae, Juncaceae, Lemnaceae, Pandanaceae, Sparganiaceae, Typhaceae, Cyperaceae,
Cyclanthaceae, s. str.
True Monocotyledones. Seeds store starch primarily in Highly Compound Starch
Grains; the vegetative organs also exhibit the same type of starch grains or other
Types and only seldom are dissolved carbohydrates found. This amylophilic group
includes Hydrocharitaceae, Pontederiaceae, Xyridaceae, Eriocaulaceae, Mayacaceae,
Araceae, Bromeliaceae, Cannaceae, Commelinaceae, Cyanastraceae, Gramineae,
Haemodoraceae, Marantaceae, Musaceae, Philydraceae, Velloziaceae, Zingiberaceae.
Derived Monocotyledones. Seeds seldom contain starch grains but storage is in form of
fatty oils and often with Reserve Cellulose. Unripe seeds, however, have Envelope
Starch Grains frequently. Vegetative organs accumulate starch plus fructosans, or
fructosans only, otherwise sucrose, glucose, fructose, or oligosaccharides. The group
is amylophobic and includes the Palmae, Agavaceae, Carludovicoideae (s. str.), Liliaceae, Alliaceae, Amaryllidaceae, Alstroemeriaceae, Taccaceae, Iridaceae, Dioscoreaceae, Orchidaceae.
NOVEMBER
1978
467
DICOTYLEDONES. The derived Monocotyledones are characterized by their accumulation
of fatty oils as reserve materials in the seeds; in the unripe seeds Envelope Starch Grains are
very often formed. This pecularity is also widespread among the seeds of the dicotyledonous
families. It is impossible to discuss the great number of these families in the present review;
it must suffice to give the numbers of families within each group. Of 191 families examined,
172 showed fatty oils in the seeds; most of these accumulated abundant small Envelope
Starch Grains in the unripe seeds. In this characteristic, the derived Dicotyledones do not
differ from the derived Monocotyledones; in this respect there is a gradual transition from
the one group to the other. The differences between the two in this regard is less than the
differences between the primary Monocotyledones and the derived Monocotyledones. The
quality of the accumulated reserve material of the seeds is not dependent on the morphologic or systematic position of the plant but only on the degree of specialization.
Some of the similarities and differences can best be shown in a table:
Primary Monocotyledones: Seeds and vegetative organs contain only Pteridophyte
Starch Grains. Except for a very few exceptions, this is a characteristic of primary
monocotyledons. However, the Haloragaceae accumulate starch grains of the Pteridophyte Type in their seeds and vegetative organs. The Nymphacaceae, s. str.,
Piperaceae, Chloranthaceae and Saururaceae store the same starch grain types as the
True Monocotyledons.
True Monocotyledones: Seeds frequently have Highly Compound Starch Grains, rarely
fatty oil (Lysichiton). Vegetative organs contain Envelope- or Envelope-Layer type
of starch grains, and fructosans (many grasses).
Derived Monocotyledones: Ripe seeds contain fatty oils, protein, plus Reserve Cellulose
or Amyloid; unripe ones have Envelope Starch Grains. Vegetative organs accumulate Envelope Starch Grains, with or without fructosans, or fructosans may occur
alone.
Primary Dicotyledones: Mature seeds frequently have Envelope Starch Grains or other
basic construction types; unripe ones are the same. Families included here are the
nine families of the Centrospermae, Polygonaceae, Cistaceae, Droseraceae, Orobanchaceae, Trapaceae, many Papilionaceae, Sterculiaceae, Fagaceae, et al.
Derived Dicotyledones: Most families belong here. Mature seeds accumulate fatty oils,
proteins, rarely Reserve Cellulose or Amyloid; unripe ones frequently have Envelope Starch Grains; vegetative organs frequently show Envelope Starch Grains or
increasingly soluble carbohydrates/Inulin.
GYMNOSPERMS. According to Engler's Syllabus II, the gymnosperms are an ancient plant
division. They are not derived from living Pteridophyta but must have developed in the
Devonian or Silurian from primitive pteridophytes, perhaps from several lines within that
group. In Wettstein (1935), the gymnosperms occupy a position between the Pteridophyta
and the Angiospermae. Examination of the vegetative organs and seeds of numerous families
in this division has yielded starch grains of the Pteridophyte Type. Undoubtedly the gymnosperms developed independently of the angiosperms; consequently, the gymnosperms are
considered in this paper after the angiosperms.
Storage forms in the gymnosperms, including a listing of the groups examined, are shown
in the following table.
Primary Gymnosperms: Amylophilic.
Order Cycadales (Cycas, Dioon, Encephalartos, Lepidozamia, Macrozamia, Zamia):
Seeds have Envelope Starch Grains. Vegetative organs (roots, leaf bases, and
stems) store large quantities of starch in the form of Envelope Starch Grains(?)
and Pteridophyte Starch Grains. Starch is stored in stems of Cycas revoluta,
Dioon edule, Macrozamia spiralis, Encephalartos caffer, Zamia spiralis, and
several other species (Samec, 1966).
Order Ginkgoales (Ginkgoaceae, Ginkgo biloba): Starch is stored in seeds as Pteridophyte Starch Grains; the same storage form is found in the stems and roots.
Derived Gymnosperms
Class Coniferophyta (order Coniferae): Seeds have fatty oils, proteins, seldom starch
but when present, Pteridophyte Type. Vegetative organs have Pteridophyte
Starch Grains in roots and stems but the stems may also store fatty oils.
468
46
TAXON VOLUME 27
4AXNVLUE2
Families examined include: Pinaceae (Abies, Cedrus, Larix, Pinus, Pseudotsuga,
Tsuga; Taxodiaceae (Cunninghamia, Cryptomeria, Metasequoia, Sciadopitys,
Sequoia, Taxodium); Cupressaceae (Callitris, Cbamaecyparis, Cupressus, Juniperus, Libocedrus, Thuja, Biota); Podocarpaceae (Phyllocladus, Podocarpus);
(Araucaria,severalspecieshavCephalotaxaceae(Cephalotaxus); Araucariaceae
Taxaceae
seeds
with
(Taxus,
starch);
Torreya).
copious
ing
Primary Chlamydospermae (Order Gnetales): Amylophilic. Welwitschiaceae (W. bainesii
only): Seeds have starch in one cell-layer plus protein. Roots have Pteridophyte
Type Starch Grains. Ephedraceae (Ephedra): Both seeds and roots and stems store
Pteridophyte Type Starch Grains abundantly. Gnetaceae (Gnetum): Abundant
starch grains in seeds; the vegetative organs have the Pteridophyte Starch Grain
Type.
Appendix: Casuarinaceae (Casuarina): Seeds store energy in fatty oil and protein; the
vegetative organs have very few Pteridophyte Starch Grains.
Summaryand Conclusions
1. Analysis of starch grains has shown that the Monocotyledones are not a
uniform class, but comprise at least three groups:
(a) PrimaryMonocotyledones. These produce starch grains in seeds and
vegetative organs and store it in the Basic Construction Type of the
Pteridophyta.The group is amylophilic.
(b) Real or True Monocotyledones. These also accumulatestarch in their
seeds and vegetative organs, but never in the form of Pteridophyte
Type Grains. The seeds accumulatemostly Highly Compound Starch
Grains, while the vegetative organs produce starch grains of other
basic types, or occasionally dissolvedcarbohydratesare formed. This
group is also amylophilic.
(c) Derived Monocotyledones. These are distinct from the other groups
in accumulating fatty oils instead of starch in their seeds. Starch is
only rarely formed in the seeds, often Reserve Cellulose. Unripe
seeds frequently accumulate Envelope Starch Grains. Vegetative organs, on the contrary, accumulatestarch and fructosans or fructosans
only, or oligosaccharides.The group is amylophobic.
2. Discovery of the peculiar Pteridophyte Type of Starch Grainin the seeds
and vegetative organs of the PrimaryMonocotyledones proves that they
could not have arisen from the Polycarpicaeor any other Dicotyledones.
3. The presence of Pteridophyte Starch Grainsin the vegetative organsof the
Pteridophytademonstratestheir mutual close affinity.
4. The Ophioglossalesare not closely related to the Marattiales.
5. Though some of the Marattiaceaehave Pteridophyte Type Starch Grains
in the rhizome parenchyma cells, the higher genera Danaea, Marattia,
Christensenia,and Angiopteris of the Angiopteridaceaeshow other starch
grain types (Envelope Layer and Nearly-LayerTypes) in the parenchyma.
These starch types are not of recent origin, for the Marattialesand Angiopteridalesare "livingfossils."
6. The same starch types (as in 5.) appear also in the vegetative organs of
some Real Monocotyledones families-Musaceae, Zingiberaceae,Marantaceae, Cannaceae,Commelinaceae,Bromeliaceae,and others, as well as the
Oxalidaceaeand other Dicotyledon families.
7. The parenchyma cells in rhizomes of Lygodium species (Schizaeaceae)
contain starch grains of the Highly-CompoundType. This type is also
found in the rhizomes and seeds of many other families-Xyridaceae,
Araceae, Gramineae (Festuceae) and in the seeds and vegetative organs of
8.
the families in the Centrospermaeof the Dicotyledones.
The starch grains in the endosperm cells of the Triticeae (Hordeae) are
especially interesting. They are formed in two steps: In young seeds, the
NOVEMBER
1978
469
grains are of the little Envelope type (0 1-6 pm); later in the ripening
seeds, these acquire a second starch cover and become larger (O
30-50 um). This is a unique occurrence in the plant kingdom.
9.
Probably all Basic Construction Types of starch grains which occur in the
cells of Monocotyledones and Dicotyledones had already evolved in the
Pteridophyta.
A full account of the data and principles summarized in the foregoing is given in
my book "Starke und Starkespeicherung bei Gefasspflanzen und einigen Vorfahren im Hinblick auf Evolution: Versuch einer Amylo-Taxonomie" (Stuttgart, Fischer, 1978).
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TAXON VOLUME 27