CONCLUSIONS
In the foregoing chapters a very limited number of plant poisons have been
considered, yet there is sufficient evidence to show that even these few
differ considerably in their action upon plant-life. This action is most
variable, and it is impossible to foretell the effect of any substance
upon vegetative growth without experiments. The degree of toxicity of the
different poisons is not the same, and also one and the same poison varies
in the intensity and nature of its action on different species of plants.
While certain compounds of copper, zinc and arsenic are exceedingly
poisonous, compounds of manganese and boron are far less deleterious,
so that a plant can withstand the presence of far more of the latter
substances than of the former. Again, the tested compounds of copper, zinc
and arsenic do not seem to stimulate growth, even when they are applied
in the smallest quantities, whereas very dilute solutions of manganese
and boron compounds decidedly increase growth. But, differentiation occurs
even in this stimulative action, for while manganese is the more effective
in stimulating barley, boric acid is far more potent for peas, the shoots
being particularly improved.
A consideration of the experimental work that has been done on this
subject of poisoning and stimulation leads one to the inevitable
conclusion that it is not true to maintain the hypothesis that _all_
inorganic plant poisons act as stimulants when they are present in very
small quantities, for while some poisons do increase plant growth under
such conditions, others fail to do so in any circumstances. It is probable
that what has been found true with the few substances tested would prove
to be similarly true over a much wider range of poisons, and at any rate
the hypothesis must be dismissed in its universal application. A more
accurate statement would be that some inorganic poisons act as stimulants
when present in small amounts, the stimulating concentrations varying both
with the poisons used and the plants on which they act.
It is quite possible for a stimulation in one respect to be correlated
with a retardation in another. In the Rothamsted experiments on the action
of manganese sulphate on barley the weaker concentrations of the salt
improved the vegetative growth, as was shown by the increase in the dry
weights, but with the same strengths of the poison the ripening of the
grain was retarded, so that, while certain of the physiological functions
were expedited, others were hindered by the action of the poison.
Thus it is evident that it is exceedingly difficult sharply to
characterise either toxic or stimulant action. In neither case is the
reaction simple--many factors may come into play and many processes are
concerned, while the effect of a so-called poison may vary in respect of
each of the functions and processes concerned. If the poison is presented
in great strength the toxic action is dominant, and probably affects
many functions in the same sense, so that the action is, so to speak,
cumulative. Lower down in the scale of concentration differentiation of
action may set in, and while some processes may still be hindered, others
may be stimulated. If the two actions balance one another an apparent
indifference may be manifested, so that it seems that such strengths of
the poison have no effect on growth, either harmful or beneficial. At
still lower concentrations, with certain plants and certain poisons, the
stimulative action overpowers the toxic effect, so that in some respect
or other improvement occurs in growth.
It is quite conceivable, however, that some poisons are truly indifferent
in weak concentrations, as no stimulation makes itself evident under any
circumstances. In these cases one is inclined to suspect that the action
is somewhat more simple, in that the toxic effects gradually diminish
until no poisonous action is manifest at very weak concentrations, and as
no stimulation is present to bring the growth above the normal with these
very weak concentrations the plant is similar to those grown without any
addition of the poison.
The _modus operandi_ of these stimulative agents is not yet fully
understood. Perhaps at the present time two main theories hold the field:
(1) that they act as catalytic agents, being valueless on their own
account, but valuable in that they aid in the procuring of essential food
substances; (2) that the stimulants themselves are of integral value for
nutrition. The French school, with Bertrand at the head, hold strongly
to the catalytic theory, maintaining that manganese and boron compounds
are able to increase growth if they are present in small quantities, as
they act as “carriers” whereby the various functions of the plant are
expedited by the increased facility with which the essential nutritive
elements are supplied. The manganese in laccase, for instance, is held
to be an oxygen carrier, whereby the oxygen is first absorbed and then
released for the benefit of the plant, the manganese being regarded as
essential for the functioning of the enzyme. But, if these elements are
_essential_, this theory seems to stop short of the truth. If certain
functions are dependent for their very occurrence upon the presence
of even minute traces of any element, then surely that element is as
essentially a nutrient element, as vital to the well-being of the plant as
is such an element as carbon or nitrogen or phosphorus, even though the
latter occurs in far greater quantity. It is necessary that one should
free one’s mind from the idea that the quantity of an element present in
a plant is an index of its value to the plant. Naturally enough, in the
early days of plant physiology, the most abundant elements first engaged
the attention of investigators, and they were divided into essential
and non-essential, ten elements being classed in the former category.
More recent work is beginning to show that other elements are constantly
present in plants, but in such small quantities that the older and cruder
methods of analysis failed to reveal them, so that until lately they have
been completely overlooked in work on plant nutrition. Even yet we do
not know which of these other elements are essential and which are merely
accidental. While we do know that the ten essential elements (C, H, O, N,
S, P, K, Mg, Fe, Ca) are necessary for the well-being of all plants, it
is conceivable that these other substances which only occur in very small
quantities may be more individual in their action, and that while a trace
of a certain element may be absolutely essential to one plant, that same
element may be quite indifferent for another species. If one takes a broad
outlook, the two theories seem to be in reality only parts of one, the
“nutrition” theory carrying matters a little farther than the “catalytic”
idea, broadening its scope and extending its application.
It seems probable that all the experimental work that has been
discussed will prove to be simply preliminary to a far greater practical
application of the principle of stimulation or increased growth. While
the physiologists have been feeling their way towards the conclusions
put forth on this subject, the agriculturists have been discovering and
extending the application of artificial manures, until at the present
time such manuring is coming into its own and is receiving more of the
widespread attention that it deserves. The possibility now exists that
in some respects the two lines of work are converging and that the
more purely scientific line will have a big contribution to make to the
strictly practical line. Artificial manuring aims at improvement of the
soil and crop by the addition of food substances that are needed in a
particular soil, a result that used to be obtainable only by the use of
the bulky farmyard manure, seaweed, &c. Apart from any other aspect of
the matter the artificials, when intelligently used, are far more easy
to handle and to regulate in supply, and they yield excellent results,
especially in conjunction with a certain proportion of organic manures.
The further prospect now opened up is the possibility of utilising some
of these stimulating compounds as artificial manures. As only small
traces are beneficial, larger amounts being poisonous, it is obvious
that only small quantities would be needed, and, as the compounds are not
usually very expensive, a considerable increase of crop for a relatively
small outlay might be anticipated if no complicating factors intervened.
Very much work will be required in the field to test the value of these
substances, as their action may be influenced by the nature of the soil,
climatic conditions, general conditions of manuring, and the crops grown.
Some tests have already been made, especially in Japan, with boron and
manganese, and these indicate a promising field for investigation.
Above all, it is most important to realise that one is approaching an
entirely unexplored field, and that it is inevitable that the results of
the initial experiments will be contradictory, at least in appearance,
so that it is necessary to keep an open mind on the subject, being ready
to modify one’s ideas as circumstances require, as improved experimental
methods lead on to more accurate results.
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INDEX OF PLANT-NAMES
_The symbols after the plant-names represent the elements referred to
on the pages indicated._
Acorus Calamus
Mn, 79
Actinostrobus pyramidalis
Mn, 80
Aesculus
Mn, 78
Agathis robusta
Mn, 80
Ageratum
Cu, 24;
Zn, 42
Alder
As, 53
Algae
As, 62, 64;
B, 66, 77
Allium
(_see_ Onion)
Zn, 47
Alnus incana
Mn, 79
Aloe
Cu, 26
Amomum sp. (Paradieskörner)
Cu, 15
Ampelopsis
Cu, 32;
Zn, 47
Anaboena
Cu, 35
Angiosperms
B, 66
Anthracnose
B, 76
Apple
B, 65, 66
Apricot
Cu, 16
Araucaria Bidwilli
Mn, 80
Cunninghamii
Mn, 80
Armeria sp.
Zn, 38
vulgaris
Zn, 36, 37
Arundo Sacchar
Mn, 78
Asparagus
Mn, 79
Aspergillus
Cu, 33, 34
Aspergillus niger (= Sterigmatocystis nigra)
Zn, 48, 49, 50;
As, 63;
B, 76;
Mn, 90
Athrotaxis selaginoides
Mn, 80
Barley, 11, 13;
Sr, 5;
Cu, 16, 17, 20, 29;
Zn, 37, 39, 40, 44, 46;
As, 52, 54, 55, 57, 60, 61;
B, 66, 69, 75;
Mn, 79, 81, 82, 83, 84, 85, 86, 89, 90, 92, 93, 94
Beans
Cu, 16, 17, 26;
Zn, 47
Beech
Mn, 79
Beetroot (Beta vulgaris)
Cu, 16, 26;
Zn, 37, 39;
Mn, 78, 79
Beet, sugar
B, 65
Betula alba
B, 66
Birch
Mn, 79
Botrytis cinerea
Mn, 90
Brassica oleracea
Mn, 78
Buckwheat, 11;
Cu, 16, 17, 29, 30;
As, 53
(_see_ Polygonum Fagopyrum)
Cabbage
Sr, 5;
As, 51, 52;
Mn, 87
Cacao
Cu, 16
Callitris gracilis
Mn, 80
robusta
Mn, 80
Cannabis
B, 72
sativa
B, 66
Capsella Bursa-pastoris
Zn, 37
Cardamomum minus
Cu, 15
Carrot
Mn, 79
Cauliflower
Mn, 78
Cherry
B, 65
Chestnut
Ca, 71;
Mn, 79
Chickpea
Cu, 16;
B, 66
Chicory
Mn, 79
Chrysanthemum
B, 66
Clover
Zn, 42, 44
Colea
Cu, 24;
Zn, 42;
As, 58
Collinsia
B, 72
Coniferae
Zn, 38
Conium maculatum
Mn, 78
Colza
B, 74
Couch grass
Mn, 83
Cow pea
Cu, 18
Cruciferae
Mn, 78, 89
Cryptomeria japonica
Mn, 88
Curcuma longa
Cu, 15
Currant
Cu, 31
Dacrydium Franklini
Mn, 80
Dianthus caryophyllus
Mn, 79
Elodea canadensis
Cu, 32;
Zn, 48
Fagopyrum esculentum
Cu, 29
Ferns
B, 66
Fig
Zn, 42;
B, 65, 66
Fir
Mn, 80
Flax
Mn, 78, 88, 89
Fungi
Cu, 33;
Zn, 44, 50;
As, 64;
B, 66, 77
Geranium
Cu, 24;
Zn, 42;
As, 58
Gramineae
B, 72;
Mn, 89
Grasses
Zn, 39, 40, 42
Gymnosperms
B, 66;
Mn, 80
Haricot
B, 71, 72
green
As, 52
white
Zn, 37
Hemp
Sr, 5
Hop
B, 66
Hordeum distichum
As, 54
vulgare
Zn, 39, 47
(_see_ Barley)
Iberis
B, 72
Laminaria saccharina
B, 66
Leguminosae
B, 72;
Mn, 78, 89
Lentil
Mn, 79
Lichen
As, 59
Linseed
Cu, 16, 17
Linum
B, 72
Love-apple
B, 66
Lucerne
Mn, 79, 91
Lupin
Cu, 17, 19;
As, 59
white
As, 61;
B, 67, 70
(_see_ Lupinus albus)
yellow
B, 70, 75
Lupinus albus
Cu, 19, 20, 22;
Zn, 45;
B, 68
(_see_ White Lupin)
Maasa picta
B, 65
Maize
Cu, 16, 17, 19, 24, 26, 27;
Zn, 37, 44;
As, 53, 54, 60;
B, 67, 68, 71, 72, 74;
Mn, 79, 81
Menyanthes trifoliata
Cu, 35
Mildew
B, 76
Molinia cærulea
Cu, 16;
Zn, 37
Mould
B, 76
Mucor mucedo
As, 59, 63
Mushroom
B, 66
Mustard
Cu, 17
Nasturtium
Cu, 17
Nuphar lutea
Cu, 35
Oak
Cu, 16;
Zn, 42
Oat
Cu, 16, 17, 19;
As, 53;
B, 74;
Mn, 79, 82, 86, 87
Onion
B, 66;
Mn, 88
Onobrychis sativa
Zn, 39
Opium
Mn, 79
Oscillatoria
Cu, 35
Panicum italicum
Cu, 26;
As, 58
Pansy
Cu, 24;
Zn, 42;
As, 58
Paprika
Cu, 16, 17
Paradieskörner (Amomum sp.)
Cu, 15
Pea
(_see_ Pisum sativum)
sweet
Cu, 17
Pear
Cu, 16;
B, 65
Penicillium
Cu, 33, 34
brevicaule
As, 63
cupricum
Cu, 34
glaucum
Cu, 23;
As, 59, 63;
Mn, 90
Phaseolus vulgaris
Cu, 17;
As, 59
Phormidium Valderianum
As, 62
Phyllocladus rhomboidalis
Mn, 80
Pine
Mn, 80
Pineapple
Mn, 83
Pisum arvense
Cu, 29
sativum
Cu, 17, 18, 26, 27, 29;
Zn, 41, 47;
As, 58
(“Pea”), 3, 11, 13, 93;
Cu, 17, 26, 29, 30;
Zn, 40, 46;
As, 51, 54, 55, 56, 58, 61;
B, 67, 73, 74, 75, 93;
Mn, 79, 81, 82, 85, 86, 87, 89, 92
Plantago lanceolata
Zn, 37
Podocarpus elata
Mn, 80
Polygonum amphibium
Cu, 35
aviculare
Zn, 37, 38
Fagopyrum
Cu, 26, 39;
Zn, 41;
As, 54, 58
(_see_ Buckwheat)
Persecaria
Cu, 5;
As, 54
Poplar
Cu, 15
Potato
Cu, 16, 26, 27, 30;
As, 52;
Mn, 78, 79
Protococcus infusionum
As, 62
sp.
As, 63
Pyrus communis
Mn, 79
Radish
Sr, 5;
B, 74;
Mn, 84, 90
Raphanus
B, 72
sativus
Zn, 39
Raspberry
As, 65
Rice
Zn, 47;
B, 66, 73;
Mn, 79, 86, 87, 88
Rosa remontana
Mn, 79
Rubus
B, 66
Rye
Cu, 16;
As, 60, 61
Sainfoin
Mn, 79
Secale cereale
Cu, 26;
Zn, 41;
As, 58
Silene inflata
Zn, 36, 37
Solanum tuberosum
Mn, 78
Soy beans
Cu, 17, 19;
B, 67;
Mn, 81, 85
Spinach, 41;
B, 73;
Mn, 89
Spergula arvensis
Zn, 39
Spirogyra
Cu, 35;
As, 62;
B, 76
Stichococcus bacillaris
As, 62
Sterigmatocystis nigra
Zn, 48, 49
(_see_ Aspergillus niger)
Sugar cane
B, 65
Taraxacum officinale
Zn, 37
Tea
Mn, 79
Thlaspi alpestre
Zn, 36
sp.
Zn, 38
Tobacco
B, 66
Trapa natans
Mn, 79
Trifolium pratense
Zn, 39
Triticum vulgare
Cu, 17;
B, 67
(_see_ Wheat)
Tropeolum Lobbianum
Cu, 17, 18
Turnip
As, 51, 52;
B, 74
swede
Mn, 78
Tussilago Farfara
Zn, 37, 38
Ulothrix tenerrima
As, 62
Ustilago
Cu, 28
Vaucheria
B, 76
Vicia Faba
Sr, 5;
Cu, 27, 29
sativa
B, 67;
Zn, 39
Viola sp.
Zn, 38
tricolor
Zn, 36
Vine
Cu, 31
Vitis vinifera
As, 52
Watercress
B, 66
Water-melon
B, 65
Wheat
Cu, 16, 17, 23;
Zn, 37, 44, 46;
As, 52, 60;
B, 66, 70, 72, 73;
Mn, 79, 81, 83, 84, 85, 86, 87
Willow
Zn, 39, 40
Yeast
Zn, 50;
B, 76
Zea japonica
Cu, 17, 18
Mays
(_see_ Maize)
GENERAL INDEX
Absorption capacity of soils for zinc, 41
of poisons by plants, 25
Accelerators, 45
Action of heavy metals in mixed solutions, 20
Adsorption, 8, 23
Aeration in water cultures, 8
Algae, assimilation of arsenic by, 62
clearing ponds of, 35
effect of arsenic on, 62
effect of boron on, 76
effect of copper on, 35
Aluminium, 45, 78
Arbutin, 4
Arsenate, potassium, 53, 62, 63
sodium, 55, 57, 61
Arsenates, 53, 57
Arsenic acid, 53, 54, 60, 61, 62, 64
acid v. arsenious acid, 53
acid v. phosphoric acid, 53, 62
elimination of, 59
gas liberated by moulds, 63
in soil, effect of, 58
in superphosphate, 58
Arsenious acid, 53, 54, 57–61, 64
immunity of plants to, 58
Arsenite, sodium, 55, 56, 61
Arsenites, 53, 57
v. arsenates, 57
Artificial oxydases, 84
soil, 24, 46
Assimilation, reduction in water plants, 48
Barium, 44
Borate, calcium, 71
potassium, 71
Borates, 72
Borax, 71, 73
Bordeaux mixture, 30
blocking of stomata by, 33
on assimilation, effect of, 33
Boric acid, 1, 65–76, 93
Boromannitic acid, 68
Boron, antiseptic action of, 70
colour due to, 75
distribution in plants, 66
poisoning, indication of, 68, 69
rôle in plant economy, 74
Cadmium, 31
Calamine, 37
plants, 38
soils, flora of, 37
Calcium carbonate, 4, 23, 25
chloride, 20, 59
sulphate, 20, 44, 45
Carbon black, 23
dust, 10
Catalytic elements, 49, 91
Chlorophyll, 44, 60
Complementary manures, 47, 91
Conditions of plant life, 7
Copper, acetate, 19
action on plant organs, 30
bicarbonate, 26
bromide, 19
chloride, 19, 20
compounds, corrosive action on plant roots, 5, 27
distribution in tissues, 16
mode of action on plants, 25
nitrate, 19, 25
oxide, 15, 25
quantity in certain plants, 17
salts, injection into plant tissue, 31
sprays, effect on leaves, 30, 32
sulphate, 5, 19, 20, 22–27, 29–35, 41
Cumarin, 4
Distilled water, preparation of, 10
Double decomposition in soil, 25
Duration of experiments, 13
Experimental methods, comparison of, 14
Ferric chloride, sublimed, 86
hydrate, 23
Fungi, effect of arsenic on, 63
effect of boron on, 76
effect of copper on, 33, 34
effect of manganese on, 90
effect of zinc on, 48
Galactose, 76
Glucose, 76
Germination, effect of arsenic on, 60
effect of boron on, 72
effect of copper on, 27, 28
effect of manganese on, 84
effect of zinc on, 43
of seeds in sawdust, 11
Grading of seeds, 11
Growth in copper-distilled water, 17
of peas in water cultures, 11
Hydrochloric acid, 22
Hydroquinone, 91
Hypothesis of universal stimulation, 93
Iodine, 2, 91
Individuality of plants, error due to, 13
of species, 61
Interaction between soil and poison, 9
Iron, 31, 49
oxide, 78
sesquioxide, 84
sulphate, 48, 81, 85, 88
Laccase, 91, 95
Lack of control over field experiments, 9
Lead, 10, 26, 42, 44
Magnesium carbonate, 83, 84, 87
chloride, 20
sulphate, 1
Manganese as top-dressing, 89
chloride, 86, 88
commercial value of, 88
cytological action of, 81, 82
dioxide, 84, 87
essential to Coniferae, 80
in Australian soils, 83
in leaves, deposition of, 82
in organic combination, 79
iodide, 83, 84
manuring, after-effects of, 88
nitrate, 83, 84
oxide, 78, 79, 83, 84, 86, 88
phosphate, 79, 83, 86
retardation of ripening by, 82, 85
sulphate, 81–89, 94
Masking effect of inorganic food salts, 4, 20
Mercuric chloride, 22
oxide, 41
Mercury, 26
Metallic oxides, 78
Methods;
field experiments, 9
sand cultures, 8
soil cultures in pots, 9
water cultures, 7, 11
Mode of entry of poisons into plants, 4
Nickel, 24, 50
Nucleinic combination, 79
Nutrient solutions, composition, 13
Oligodynamic action, 28
Over-mineralisation of plants, 71
Phosphoric acid, 53, 54, 62, 64
Photo-synthesis, effect of copper on, 32
Potassium hydrate, 22
Presence of arsenic in animals, 51
in plants, 51
of boron in plants, 65
of copper in plants, 15
of manganese in plants, 78
of zinc in plants, 36
Pyrogallol, 91
Raulin’s solution, 49
Reproduction of poisoned plants, 40
Silver nitrate, 22
Sodium chloride, 20, 44
hydroxide, 22
nitrate, 4
Sterile cultures, 24
Stimulation, by injection of copper solutions, 31
by small doses of poisons, 2
definition of, 2
local, 47
of Aspergillus niger, 50
of fungi by copper, 34
of plants by arsenic, 61
of plants by boron, 73
of plants by copper (negative), 28
of plants by manganese, 84
of plants by zinc, 45–47
physiological considerations of manganese, 90
Strontium sulphate, 5
Sugar, 22, 31, 48, 49, 50, 68
Sulphur, flowers of, 10
Thymol, 22
Toxic action, effect of arsenic, 52
effect of boron, 67
effect of copper, 17
effect of light on, 44
effect of manganese, 81
effect of zinc, 38
equivalent, 18
limits of plant poisons, estimation of, 26
Toxicity, of nutrient salts, 1
of organic compounds, 4
of poisons, cause of, 22
of positive ions in copper compounds, 19, 22
reduction of, 39, 44
reduction of, by carbon black and ferric hydrate, 23
reduction of, by insoluble substances, 22
Toxin and nutrient, distinction between, 3
Transmission of power of resistance, 72
Valency, effect on toxicity, 44
Vanillin, 4
Variation in results on different substrata, 24
Zinc, absorption by roots, 42
carbonate, 38, 42, 43
effect of, on lower plants, 48
effect of, on plant and soil, 41
fixation of, 49
mode of action on plants, 43
oxide, 37, 47
oxide on leaves, deposition of, 47
storage in seeds, 43
sulphate, 38–49
sulphide, 42
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Transcriber’s Notes
Minor inconsistencies in punctuation have been standardised--in
particular, missing periods and parentheses have been supplied where
obviously required. All other original errors and inconsistencies have
been retained, except as follows:
Page 27: changed Just to Just’s
(in Just’s _Bot. Jahresber._)
Page 37: Added missing decimal point
(total ash, (·006% of the)
Page 76: changed gms. to gm.
(in until 5 gm. boric)
Page 97: changed in to im
(Vorkommen von Iod im Thierkörper.)
Page 98: changed and to et
(Colin, H. et Lavison, J.)
Page 100: changed Experiments to experiments
(Pot experiments to determine the)
Page 101: changed cœrulea to cærulea
(Zusammensetzung von Molinia cærulea)
Page 101: changed and to und
(Hiltner, L. und Gentner, G.)
Page 101: changed Waar. to War.
(Nahrungsmittel Hyg. War. IX, 1–4.)
Page 102: changed Genuss-mittel to Genusmittel
(Nahrungs- und Genussmittel,)
Page 102: added date to entry
(IV, 489–92. (1901))
Page 102: changed Rübensache to Rübenasche
(Bestandtheile der Rübenasche. Ber.)
Page 103: changed Magenesium-salze to Magnesiumsalze
(Calcium- und Magnesiumsalze im)
Page 105: changed Vorhandsein to Vorhandensein
(das Vorhandensein von Arsen)
Page 105: changed Pflanzen-production to Pflanzenproduction
(der Pflanzenproduction. Casopis pro)
Page 105: changed prumsyl to prumysl
(Casopis pro prumysl chemicky,)
Page 107: added entry for Cu under Fungi
(Cu, 33;)
Page 110: changed Photosynthesis to Photo-synthesis
(Photo-synthesis, effect of copper on, 32)