A doyen of Indian botanists: H. Y. Mohan Ram

LIVING LEGENDS IN INDIAN SCIENCE

A doyen of Indian botanists: H. Y. Mohan Ram

Ramesh Maheshwari, K. R. Shivanna, R. Dore Swamy, K. Sankara Rao and Gita Mathur

Holenarasipur Yoganarasimham Mohan the beginning of our own scientific feet; age 2200 years; and weight 1256
Ram, fondly referred to as HYM (Figure careers, we suppose HYM would take metric tonnes. Seeing ‘General Sher-
1), belongs to the rare category of gene- umbrage at being labelled or categorized man’, the questions that came to mind
ral botanists that is rapidly becoming as a specialist. In his own words, he has were the methods used to determine the
extinct. He has completed 83 years. On been a teacher, researcher, naturalist, weight of a living tree.
this occasion, we offer this academic author, editor, reviewer, adviser, pro-
tribute, sautéed with personal anecdotes moter and popularizer of science, travel- Making of a ge neral botanist
for transmitting to us his passion for ler, photographer, lover of music and
plants in general, and of the flora of cricket and a tolerably good cook. He After his postgraduate training under M.
Amazon and of Malabar in Kerala, in would always find the time to go out of Anantaswamy Rau in Mysore, HYM was
particular. We briefly summarize his the way to look up his former students or attracted to take up research on plant
major research contributions on flower colleagues to popularize botany, espe- embryology, which had then begun to
colour, sex expression in flowering cially economic botany of trees valued as emerge as an important discipline due to
plants, and in vitro culture of bamboo sources of timber, gums and oils. For ex- publication in 1950 of a book titled An
and aquatic angiosperms. Additionally, ample, in a recent article on sandalwood Introduction to the Embryology of Angio-
we sum up what each of us has learnt he covers the use of the fragrant heart- sperms by PM. HYM researched on the
from him over informal meetings when- wood in carving, the use of its oil in per- seed development in some plants belong-
ever he visits Bangalore and reflect on fumery and toiletry and in combating ing to the family Acanthaceae. After ob-
our perceptions of HYM as a general antibiotic-resistant bacteria, and points taining his Ph D, on PM’s advice, HYM
botanist and a humanist. out the need for research on phytoplasma went to the laboratory of F. C. Steward
causing spike disease, micropropagation (see Figure 1, centre) at Cornell Univer-
As general botanists do not come any of sandalwood, and on the selection of sity, who was attracting international
more, it is of interest to know how HYM diploid and triploid somatic embryos for attention for his contributions to the con-
became one. Born in Karnataka, HYM establishing superior genotypes having cept of totipotency. Using the tissue cul-
obtained his early education in Mysore. higher quantity of heartwood. ture technique by which excised tissues
After obtaining his Master’s degree in are grown in vitro in artificial nutrient
botany, he went to Agra for research on At the time of writing this article, one media, Steward, trained as a chemist,
the development of seeds under the guid- of us (K.R.S.) was visiting California demonstrated that plant cells are toti-
ance of P. Maheshwari (PM), who was where, inspired by HYM’s approach of potent – that is, each plant cell has the
then at Agra. Subsequently, when PM learning based on personal observations genetic information for developing into a
moved to University of Delhi, HYM was of flora in nature, he visited the Pacific complete plant. For example, cells from
appointed a lecturer in Delhi University West Coast to see the endemic redwood
from where he retired as a professor. trees, named for the colour of their bark
Currently, HYM is a research professor and heartwood. One specimen of Sequo-
of the Indian National Science Academy, iadendron giganteum (Figure 2), named
working at Shriram Institute for Indus- ‘General Sherman’, has a diameter 36.5
trial Research, Delhi. Knowing him since feet; circumference 103 feet; height 275

Figure 1. H. Y. Mohan Ram (right side, photo by Gita Mathur) in group photo taken in Figure 2. ‘General Sherman’ (Sequoia-
Botany Department at Delhi University. This photograph taken during an International dendron giganteum) growing in California
Symposium held in 1959 on Plant Tissue and Cell Culture shows, sitting from left to is one of the largest known trees in the
right: N. S. Rangaswamy (Convenor), B. M. Johri, J. Reinert, F. C. Steward, P. world. Photo by K. R. Shivanna.
Maheshwari, J. Swarbrick, H. E. Street, J. P. Nitsch and B. Sen. H. Y. Mohan Ram is
standing at the extreme left in the last row. 305

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LIVING LEGENDS IN INDIAN SCIENCE

the phloem tissue of carrot roots cultured and bladders with tiny hair-like projec- virus-free (bunchy top, bract mosaic, in-
in suitable sterile nutrient media differ- tions at their opening. Commonly known fectious chlorosis, streak) shoot tips
entiated into plantlets that could be trans- as the bladderwort, its scientific name is through plant tissue culture and derived
ferred to soil and grown into complete Utricularia. The bladders trap tiny in- plantlets from meristem culture of the
plants, thereby demonstrating that each sects. As it generally grows in nitrogen- high-yielding varieties are mass propa-
cell of the adult plant retains the genetic deficient habitat, it was thought that this gated in the field (Figure 3). In vitro
potential to regenerate an entire plant. plant depends on insect-derived nitrogen propagation of banana is the most suc-
HYM’s exposure to tissue culture tech- for its growth. One of us (R.D.S.) took cessful commercial achievement in India.
nology led him to initiate work in this up this problem for Ph D. By raising
field on his return to India. He guided his seed-cultures of U. inflexa in a synthetic Sex expression in Cannabis
Ph D students to apply this technique to nutrient medium containing only an
understand many aspects of plant growth inorganic nitrogen source, it was demon- Cannabis sativa (hemp or marijuana) is
development (Figure 3). With his train- strated that Utricularia can utilize inor- dioecious, comprising male and female
ing in systematic and experimental bot- ganic nitrogen for its growth. However, plants, but sex becomes distinguishable
any, his own curiosity and self-education the in vitro grown plants flowered only only when the flowers are formed. The
through observations and enquiries, when the cultures incubated under short term ‘marijuana’ refers to its medicinal,
HYM established himself as an effective day conditions (20 cycles of 16 h dark recreational or spiritual use involving the
teacher and a researcher. Most impor- and 8 h light). This was the first report of smoking of its flowers. Reportedly male
tantly, his broad outlook on plants and in vitro culture of an insectivorous plant. and female flower buds vary in hallu-
his expertise in identifying them and We became impressed with plant organ cinogenic effect. Smoking leaves are
pointing out their economic uses attracts and tissue culture technique for demon- used for relaxation, stress relief, and for
both botanists and the laypersons. HYM strating the regenerative power of plants. an overall sense of calm and serenity for
has been a popular plant scientist; and is The technique allows plant cells and body pain relief and in the treatment of
invited for giving lectures and as an ex- tissues to be indefinitely maintained in insomnia. As female plants contain about
pert in the selection and appointment of culture by their successive transfers to 5–10% or more delta 9-tetrahydrocanna-
new cadres of staff and faculty in teach- fresh medium at regular intervals of time binol (THC), it was of significance that
ing and research institutions throughout and to study their physiological require- one of HYM’s students, V. S. Jaiswal,
India. From years of observations of ments for growth and reproduction. could convert the male plants into
plant diversity and their adaptation to females by spray application of ethephon
habitats, he has become a natural author- Banana tissue culture (2-chloroethanephosphonic acid), a growth
ity, commanding the respect of people regulator. Stopping the application of
like only a few can. With his cool tem- In 1958, HYM received a Fulbright ethephon caused the plants to revert to
perament and power of persuasion and scholarship for advanced research in their original sex. They hypothesized that
the desire to help move the botanical Steward’s laboratory at Cornell Univer- in Cannabis, gibberellic acid (GA) and
research forward, HYM has been an sity. Steward was interested in the bio- ethylene act as male and female hormo-
automatic choice for presiding over na- chemistry of the banana plant. Sponsored nes respectively, and that the expression
tional meetings, and in the process meet- by a research grant from the United Fruit of sex is controlled by a balance between
ing people, and updating knowledge and Company, while in Steward’s laboratory, their endogenous levels. Abscisic acid
expanding his influence on development HYM initiated work on tissue culture of (ABA) is able to overcome the GA-
of botanical science in India. the banana varieties for studying the induced male flower formation. Report-
differences in their response to nutrient edly, male or female plant flower buds
Highlights of some contributions media containing different plant growth vary in hallucinogenic effect. Fertile
regulators. The experiments on banana female flowers were induced in male
Insectivorous plant involved culturing the white mass of plants by ethephon (2-chloroethanephos-
cells or a triploid edible tissue or the phonic acid) and NIA 10637 (ethylhy-
An insect-feeding plant found floating in pulp which presumably is the endosperm drogen-l-propylphosphonate).
water bodies has fine thread-like leaves and deriving a mass of callus cells that
could be regenerated into plantlets by in- Bamboo
Figure 3. Multiplication of banana by corporating plant growth regulators, such
shoot tip culture. Photo by R. Dore Swamy. 2,4-dichlorophenoxyacetic acid or 2,4- On the basis of flowering, the bamboos
trichlorophenoxyacetic, or the supple- have been categorized under three broad
mentation of nutrient growth medium groups: (i) those that flower annually or
with coconut water that contains a com- nearly so (Indocalamus wightianus,
plex of compounds. However, these were Bambusa atra, Ochlandra sivagiriana),
ineffective. The important finding was (ii) those that flower sporadically or
that tissue from unripe fruits could be in- irregularly (Chimonobambusa sp., Arun-
duced to callus but which failed to dif- dinaria falcata, Dendrocalamus gigan-
ferentiate into plantlets. T issue from ripe teus, D. hamiltonii, D. longispathus), and
fruits did not respond. However, today (iii) those that flower gregariously once
banana plants are being cloned from the in their life in a synchronized manner

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LIVING LEGENDS IN INDIAN SCIENCE

producing enormous amount of seeds and brings about ‘thingtam’ famine. Conven- dents, Gita Mathur (GM) noted that their
then die (monocarpic) (Bambusa bambos, tional propagation of bamboos involves colour changes to orange, scarlet and
B. polymorpha, B. tuldatundra, Ochlandra propagules such as rhizomes, suckers magenta as the flowers age (Figure 4)1.
travencorica, Dendrocalamus strictus). and culm cuttings. Because a large num- By suspending coloured sticking screens
The interval between flowering seasons ber of plants can be raised with minimum on lantana bushes, GM and HYM
of a species is highly variable. For ex- effort, propagation by seed is the most showed that thrips were attracted only to
ample, Phyllostachys bambusoides takes convenient method; but seeds are not yellow colour. A methanol extract of
120 years to flower. A majority of bam- available regularly. As tissue culture Lantana pollen was also able to simulate
boos belong to the third category. We technology offers special advantages for the effect of pollination, suggesting the
know very little about how timing of bamboo propagation, HYM’s group involvement of a pollen factor2. The
flowering is programmed into a given made the first successful report on post-pollination shift in petal colouration
species of bamboo. With the availability somatic embryogenesis and plantlet re- is caused by the masking of carotenoids
of large amount of nutritious seeds avail- generation from the seed cultures of B. by differential amounts of anthocyanin.
able in the flowering year, there is rise in bambos and D. strictus via a callus. Re- GM and HYM proposed that colour
the populations of squirrels, birds and producible procedures for rhizome initia- changes may play a role in conserving
rodents. After consuming the bonanza of tion and acclimation to potted conditions pollinator energy and on the production
bamboo seeds, the populations of rats were developed. Over 10,000 plantlets of nectar by the flowers.
turn to the standing crops and cause raised by tissue culture have been intro-
severe devastation, resulting in famine. duced in various parts of India. Flora of Karnataka
In India, Mizoram has often experienced
mass seeding of bamboos and resultant Post-pollination changes in flower HYM often spoke of India’s rich floral
famines. Tribal lore and forest records colour wealth, its diversity and discussed criti-
indicate a regular flowering cycle of cal issues related to conservation. He is
about 48 years for Bambusa tulda and The newly opened flowers of Lantana also appreciative of any work on infor-
Melocanna baccifera in Mizoram. Flow- camara are rich in -carotene and are mation building on India’s flora. One of
ering of M. baccifera causes a famine yellow-coloured. One of HYM’s stu- us (K.S.R.) is developing a database of
called ‘mautam’ and that of B. tulda about 4000 angiosperms and ferns from
Karnataka based on the voucher speci-
Figure 4. Colour change in flowers of Lantana camara following pollination. Flower mens in the Herbarium JCB at the Centre
development was divided into six stages. The data was statistically analysed and is for Ecological Sciences (CES), Indian
graphically depicted. Photo by Gita Mathur. Institute of Science, Bangalore to gener-
ate a readily accessible information sys-
CURRENT SCIENCE, VOL. 106, NO. 2, 25 JANUARY 2014 tem to help evolve strategies of effective
conservation and sustainable utilization
of plant resources in the state. Herbarium
JCB was built by Cecil J. Saldanha and
subsequently transferred to CES. It
houses about 14,500 specimens belong-
ing to more than 4000 species of angio-
sperms, ferns and lichens. The duplicates
have been deposited with herbaria of
Royal Botanic Gardens at Kew, England
and the Smithsonian Institution, Wash-
ington DC, USA. The taxonomic data
presented on each species in the database
include information presented on the
herbarium specimen label, namely species
name, author citation, sub-species if any,
variety if any, family, subfamily, collec-
tion number, location, date of collection,
habitat and the collector’s name.
Scanned herbarium specimens and the
image gallery (‘synthetic flora’) allow
quick identification of species. Areas
stripped-off their native plants can be re-
stored by transplantation. The database
could be a metric of the rate of loss of
the species likely to go extinct. Figure 5
shows a scanned image of herbarium
specimen of Bruguiera gymnorrhiza of

307

LIVING LEGENDS IN INDIAN SCIENCE

Rhizophoraceae endemic to Karnataka. Delhi developed an in vitro technique to treasure trove contained Psilotum (Figure
The flowers are solitary, axillary and germinate the seeds of Podostemaceae in 7) – a plant spotted between moist rocks.
pink in colour. The seed germinates liquid medium, enabling a detailed histo- Psilotum is regarded as a primitive plant:
while the fruit is still attached to the tree logical study of the origin and develop- it has dichotomous green branches, but
(vivipary). ment of the thallus and induction of lacks leaves, roots, flowers, fruits and
flowering. They have also carried out seeds. The branches bear sporangia at
Biology of Podostemaceae field and laboratory studies to understand their tips, and the plant reproduces by
the structure of the thallus, floral bio- spores formed in the sporangia. Psilotum
The family Podostemaceae (riverweed logy, mechanism of pollination and seed is reported to form a relationship with a
family) is little studied. The aquatic production. In Indotristicha ramosissima fungus known as mycorrhiza in its envi-
plants are found in flowing waters adher- (Tristichoideae), an ephemeral primary ronment. The fungal hyphae ramify in
ing to rocks, although the mechanism of shoot axis is formed apically between the the environment, and absorb and transfer
their adhesion in the water remains un- cotyledons. No tap root is formed, but the mineral nutrients to the plant. Both
known. Three of HYM’s Ph D students two or three adventitious roots develop at Psilotum and the mycorrhiza survive in
showed that adult plants are quite unlike the radicular pole, which become com- severe nutrient-limited conditions. We
the flowering plants, lacking clearly de- pressed and grow horizontally, adhering remember this excursion in particular as
fined roots and stems with a thalloid to the stones and form flowers and fruits NSR, who has a knack for naming,
plant body resembling an alga, lichen or on vertical shoots (Figure 6), which dubbed the excursion as the ‘Fresh and
a liverwort. Nonetheless, they are flow- dehisce to liberate tiny seeds. The seeds the best’ after the name of the lodge
ering plants (angiosperms) that form lie on rocks until the next monsoon and where we were put up for the night. In
seeds. The plant body has been inter- germinate with the first showers. HYM’s this excursion it was firmly driven home
preted as root, stem or a combined shoot students have outlined the biological that making keen observations in nature
(leaf–stem) structure. This confusion has challenges that the Podostemaceae offer not only sharpens one’s eyes to catch
mainly arisen due to difficulty in study- and hope that plant scientists would use minute details, but also imparts intellec-
ing the development of the seedling. them as experimental systems. tual thrill.
HYM’s group at the University of
A feel for classical botany An example of HYM’s feel and con-
cern for plants and for classical botany is
We wish to touch upon an inspiring typi- reflected in his writings. Until recently,
cal botanical excursion during the under- when during one of his visits to Kerala
graduate years of one of us (R.M.) he met K. S. Manilal3, HYM was not
conducted jointly by HYM and N. S. aware of his work. Devoting over 35
Rangaswamy (NSR). It was an excursion years of intense labour, Manilal has
to Pachmarhi (Madhya Pradesh). Dusk translated the entire text of Hortus mala-
had befallen and several plant specimens baricus (Malabar Garden) running into
that we had collected and saved in a vas- 12 volumes – the most comprehensive
culum during our trekking remained to printed work on plants of the Malabar
be identified. With the Floras and a dis- coast that stretches about 900 km in
secting microscope unpacked, and all of length and varying from 74 to 200 km in
us holding flashlights for additional width from Goa to Kanyakumari and
illumination, HYM and NSR began dis- published between 1678 and 1693 in
secting the flowers from the specimens Malyalam on palm leaf by Itty Achuden,
and identifying them based on floral a priest–physician. HYM writes, by
characteristics. The work continued until translating and interpreting this valuable
late night, unmindful of the dinner. Our document in English, Manilal has con-
tributed significantly towards the safe-
guarding of our natural plant wealth and

Figure 5. An image of a herbarium Figure 6. Zeylanidium lichenoides, a Podostemaceae growing on a rock in Kerala.
specimen of Bruguiera gymnorrhiza (L.) Photo courtesy of Rajesh Tandon.
Lam of Rhizophoraceae. It is a small-
sized tree with tough leaves, stilt roots
and endemic to Karnataka. The flowers
are solitary, axillary and pink in colour.
The seeds germinate while the fruits are
still attached to the tree (viviparous ger-
mination). Based on some 4000 scanned
images, database of the Flora of Karna-
taka is being developed. Photo by K.
Sankara Rao.

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LIVING LEGENDS IN INDIAN SCIENCE

You are my trainer and well wisher
And your polishing reflects in all I do.

I always think of you when I need
strength

Hearing your voice generates energy in
me

The feel your blessing hand on my head
Makes me think positively, whatever the

crisis is.

Figure 7. Psilotum, an early (primitive) land plant transplanted in soil. Note sporan- I thank you Guru for Your affection and
gium at the axils of branches. Photo by K. Sankara Rao. Your tireless hours of hard work
To make me the person I am today and
indigenous knowledge. We are impressed means of finding out what plant species For giving vision to see brightness in the
that during one of our meets, HYM emo- have been lost in 300 years since van
tionally briefed us on Manilal’s collec- Rheede and get some idea on their rate of dark.
tion and identification of 996 species of extinction.
flowering plants belonging to 559 genera 1. Mohan Ram, H. Y. and Sujata Varadara-
and 134 families. These included rare In the above paragraphs, we have re- jan, Resonance, 2009, 14, 1003–1017.
and endangered endemics, wild relatives flected on our perceptions on how keen
of black pepper, cardamom, ginger and observations of plants and asking ques- 2. Mathur, G. and Mohan Ram, H. Y., Phy-
turmeric, the Malabar daffodil orchid tions about their strategies of growth and tomorphology, 1986, 36, 79–100.
(Ipsea malabarica discovered in 1850 survival have made HYM a general
and never seen again), 12 orchids that botanists rather than a specialist. Asked 3. Mohan Ram, H. Y., Curr. Sci., 2005, 89,
occur only in a pollution-free environ- what his views are on classical botany 1672–1680.
ment and 7 species new to science. being displaced by molecular botany,
Manilal and his associates have pub- HYM referred us to his views4: ‘There 4. Mohan Ram, H. Y., J. Biosci., 2002, 27,
lished over 40 research papers on their has been an over-emphasis on areas such 651–664.
explorations in this region and a refer- as molecular biology, biotechnology and
ence book, Flora of Silent Valley Tropi- genetic engineering. To assume that ACKNOWLEDGEMENTS. We thank N. S.
cal Rain Forests of India. Manilal’s work these subjects would replace the other Rangaswamy (formerly at Delhi University)
sends a clear message to those scientists well-established branches of biology and Anantanarayanan Raman (Charles Sturt
who tend to avoid research that demands appears illogical as they are all intercon- University, Canberra) for suggestions that
rigorous and intensive labour. There is a nected. It has been said that even the helped improve the manuscript.
general impression that meaningful long- most traditional branches of biology –
term studies spanning several decades systematic, anatomy, embryology and RAMESH MAHESHWARI1,*
cannot be successfully undertaken by physiology – are still needed not just as K. R. SHIVANNA2
individuals. HYM writes, ‘Even scien- data but also because all of them are end- R. DORE SWAMY3
tific institutions are reluctant to do so; less unfinished frontiers and all of them
that Manilal has accomplished this gi- are still needed to round our view of the K. SANKARA RAO4
gantic task with practically no financial living world. Each discipline seems to GITA MATHUR5
support from any government or private have a golden period, and many of them
agency deserves deep appreciation’. have several. But even after the law of 1Formerly at Department of
HYM concluded with the plea that the diminishing returns has taken over, there Biochemistry,
scientific communities recognize Manilal’s is no justification for abolishing a disci-
scholarly and painstaking achievement at pline that has become “classical” ’. Indian Institute of Science,
a time when citation indices and working Bangalore 560 012, India
in high-tech areas using modern tools HYM is a botanist who has ploughed 2Formerly at Department of Botany,
appear to be the principal criteria for areas of enquiry untouched by others. In University of Delhi,
judging merit. Even if Manilal’s volumes closing, one of us (GM) would like to Delhi 110 007, India
may not be in our reading or reference dedicate this poem to her guru, HYM: 3Formerly at Indian Institute of
list, acquaintance with this work offers a
My teacher, guide and supervisor Horticultural Research,
Are not sufficient when I think about you Bangalore 560 089, India
4Centre for Ecological Sciences,
Indian Institute of Science,
Bangalore 560 012, India
5Department of Botany,
Gargi College,
Delhi 110 049, India
*e-mail: ramesh.maheshwari01@
gmail.com

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