Wednesday, December 30, 2015

The last Christmas truce

A modified version of this post was published at The wire: 


It was December 1914. The rain had finally relented after days. The ground was mired in trenches flooded with water. There was continuous sniping, machine-gun fire and artillery shelling which had left a deathly trail of casualties behind.

“It was a Christmas card Christmas eve. There was white beautiful moonlight, frost on the ground – almost white everywhere. And round about – I should think – 7 or 8 in the evening we heard this singing and a lot of commotion and we saw some lights.”
“They finished their carol – we applauded them then we thought we must retaliate in some way so we replied with ‘The First Noel’.”
“So we went on…. Well, I thought this was rather an extraordinary thing really – to think of the two nations both singing the same carol in the middle of a war.”

Yes, this did happen. An unofficial ‘Christmas truce’ did happen in the middle of The First World War in December 1914. The first war of its kind was also marked by the last truce of its kind.

‘The Christmas Day Truce of 1914,’ a lithograph by Arthur C. Michael published on Jan. 9, 1915, shows British and German soldiers out of the trenches of World War I, arm in arm and exchanging headgear. ARTHUR C. MICHAEL/THE ILLUSTRATED LONDON NEWS PICTURE LIBRARY, LONDON, UK/BRIDGEMAN IMAGES

Declaration of the war

A hundred and one years ago, the world was unexpectedly plunged into the First World War. Europe was in the throes of great economic prosperity with a young German nation leading the way. In his very popular book at the time - The Great Illusion, Norman Angell had argued that war between industrial countries was futile and that economic interdependence between industrial countries meant that a war would be economically harmful to all countries involved. This was seen as a big deterrent to any possible threat of war. And yet, spurred by a complicated system of secret ententes and alliances, the allied countries – United Kingdom, France and Russia had gone to war against the Central powers of Germany and Austria-Hungary in July 1914 over the tragic assassination of Prince Archduke Franz Ferdinand and Princess Sophie of Austria-Hungary.

It was the first global confrontation of alliances that went beyond nations, and even continents, to involve the whole world through direct or indirect interests. The war had begun in the summer of 1914 and people were certain that the “troops will be home when the leaves fall”. Instead, what followed were four years of mass slaughter in deep, muddy trenches with barbed wires, battle tanks, explosives, aerial bombing, and chemical warfare that led to an entire generation of missing men.

Upon the declaration of war in July, a regimented and extra-ordinarily disciplined German advance had routed through Belgium despite stiff resistance and arrived at the gates of Paris only to be checked by the British Expeditionary Forces (BEF) at the last instant. After five months of industrial scale slaughter, the war came to a standstill on the western front in November 1914 with opposing armies staring at each other from across hastily built trenches that flanked the no-man’s land.

Life in the trenches

Life in trenches was no fun at any time of the year with the continuous dread of gunfire and shelling, flooding due to rains, and the poor hygienic conditions that arose from living in them amidst the wounded and the dead; but December was especially bad with its wet, frigid conditions. Early in December 1914, Pope Benedict XV had proposed an official “truce of god” that would cease fighting over the Christmas period but the very idea of a truce was rejected by the powers that be on both sides.

                              


Life in the trenches, Image from Chris Baker’s book: The Truce – the day the war stopped (http://www.1914-1918.net/truce.htm)





However, in order to maintain morale of the troops, throughout the month of December, 460,000 parcels and 2.5 million letters were sent to British soldiers in France. King George V in UK sent a card to every soldier and his daughter, Princess Mary, lent her name to a fund that sent a box of gifts (filled with chocolates, butterscotch, cigarettes, tobacco, her picture and a facsimile of George V’s greeting to the troops - ‘May God protect you and bring you safe home’) to all serving soldiers. The Germans at their end, sent tabletop Christmas trees (Tannenbaums), tobacco/cigars and festive wreaths for the soldiers to celebrate. Clear skies and a brief reprieve from the rain further lifted the spirits on both sides as a cold frost settled in, setting the stage for a white Christmas. It was in this context that the unprecedented ‘Christmas truce’ happened.

The truce

Many differing oral accounts, diary entries and letters suggest that the truce emerged spontaneously on the battlefields of the Western Front; and yet, despite these many accounts to this day no one knows exactly where it began or how it spread through the trenches. In all, about 100,000 people are believed to have participated in the spontaneous Christmas Truce of 1914.

On Christmas eve, candles and trees went up along parts of the German frontline. It is believed that they also delivered a chocolate cake to the British line accompanied by a note that proposed a ceasefire so that the Germans could have a concert. British officers at the frontline accepted the proposal and offered some tobacco as their present to the Germans. The goodwill soon spread along the 27-mile front line as carols echoed one after the other alternating between the German and British camps.





Encouraged by the events of the night before, on Christmas morning, German soldiers are believed to have emerged from their camps, calling out “Merry Christmas” in English. Frank Richards, a British soldier who experienced the “Christmas Truce” says, “On Christmas morning we stuck up a board with ‘A merry Christmas’ on it. The enemy has stuck up a similar one….. Two of our men then threw their equipment off and jumped on the parapet with their hands above their heads. Two of the Germans done the same and commenced to walk up the riverbank, our two men going to meet them. They met and shook hands and then we all got out of the trench. Buffalo Bill [the Company Commander] rushed into the trench and endeavored to prevent it, but he was too late: the whole of the Company were now out, and so were the Germans. He had to accept the situation, so soon he and the other company officers climbed out too. We and the Germans met in the middle of no-man's-land. Their officers was also now out. Our officers exchanged greetings with them.”

What began as general bonhomie and exchange of greetings led to the exchange of cigarettes, chocolates, cognac, rum, food, buttons and other wartime supplies. There are also reports of many small-scale football kick-abouts and other communal activities like pig-roasts. While the soldiers exchanged gifts and took photos spontaneously, it was also an opportunity for them to leave the squalid trenches and to tend to the wounded and dead in the No man’s land. The troops from both sides could also finally bury their dead comrades whose bodies had lain for weeks on end. In many places, the truce extended well into boxing day (the day after Christmas) and each side seemed to wait for the other to initiate firing. In fact, Murdoch M. Wood, a British soldier is reported to have said: “I then came to the conclusion that I have held firmly ever since, that if we had been left to ourselves there would never have been another shot fired.” It was however only a truce and hostilities eventually returned; in some places sooner than in others.

The truce was widespread but not universal as the firing is reported to have continued in many sectors like at Yser where bloody battles took place over Christmas. In fact, the French troops are reported to be puzzled and peeved by these reports of the British troops hobnobbing with their German enemies. Not surprisingly, young Adolf Hitler, then a Corporal of the 16th Bavarians, shared a similar opinion himself and said: “Such a thing should not happen in wartime. Have you no German sense of honor?”

The anger from the top

As reports of the truce reached the generals, the High Command was angry as it feared a mutiny due to fraternization with the enemy. Many commanders believed that this proximity and fraternization between the troops posed “the greatest danger” to the morale of soldiers and told divisional commanders to explicitly prohibit any “friendly intercourse with the enemy”. General Smith-Dorrien issued an instruction to commanders of all Divisions: "It is during this period that the greatest danger to the morale of troops exists. Experience of this and of every other war proves undoubtedly that troops in trenches in close proximity to the enemy slide very easily, if permitted to do so, into a "live and let live" theory of life...officers and men sink into a military lethargy from which it is difficult to arouse them when the moment for great sacrifices again arises...the attitude of our troops can be readily understood and to a certain extent commands sympathy...such an attitude is however most dangerous for it discourages initiative in commanders and destroys the offensive spirit in all ranks...the Corps Commander therefore directs Divisional Commanders to impress on subordinate commanders the absolute necessity of encouraging offensive spirit... friendly intercourse with the enemy, unofficial armistices, however tempting and amusing they may be, are absolutely prohibited". Strict orders were thus issued to end any such interactions between the troops along with harsh punishment for any man who refused to fight. And these policies ensured that such a widespread cessation of hostilities was never again seen in the war.


General Sir Horace Smith-Dorrien – commander of British 2nd Army Corps Expeditionary Force – issued strict warnings to his senior officers about preventing fraternization with enemy soldiers.
  
‘Live and let live’

The Christmas truce, however, was not unique as the massive violence of the war had indeed engendered an ethos of “live and let live” between warring troops. Infantries in close proximity would engage in friendly banter and barter of goods. In some areas, there was also an implicit agreement to not shoot at men while retrieving their dead/injured, while exercising or during meal times. Wagon trains delivering food behind enemy lines were easy targets for artillery but destroying the enemy’s food would elicit the same reaction from them; thus food wagons were mostly left unharmed. The same process made latrines safe zones. Such truces emerged repeatedly during the war and the ‘army top brass’ in the rear would intervene by rotating troops, threatening courts-martial and ordering savage raids that required hand-to-hand combat. There have also been other instances of occasional ceasefires between troops in the war but the Christmas truce of 1914 is unique for its sheer size, scale and spontaneity.

Even today, more than a century after the truce, it is remembered as a testament to the power of humanity and of the individual soldier on the frontlines. Its legend has been memorialized through books, movies and advertisements. To mark the centenary last year, Prince William unveiled a memorial with a metal frame representing a soccer ball with two hands clasped inside. The truce provided an unforgettable memory for many such as the British soldier who confessed in a letter the following day, “I wouldn’t have missed the experience of yesterday for the most gorgeous Christmas dinner in England.”

In the middle of an unexpectedly long and disastrous conflict, the truce symbolizes our desire for peace and our ability to effect change from the grassroots, no matter how fleeting. Instead of an organized, top-down ceasefire, the truce of 1914 was a spontaneous series of armistices that originated from the men in the trenches. Soldiers on the battlefront reclaimed the peace that was denied to them by their governments and generals as a matter of policy. As Alfred Anderson from the Fifth Battallion, the Black Watch later told: “It was a short peace in a terrible war.” But sometimes that is all that you need – a cessation of hostilities to see the enemy and his human side. It may not give you everything you want but it leaves a lasting impression for sure.

Sources:

Wednesday, December 31, 2014

The flutter of a butterfly…

The butterfly effect is a common trope in popular science and fiction that attempts to convey a hard-to-grasp scientific fact: a very small change in the initial conditions can create a significantly different outcome. The phrase refers to the idea that a butterfly's wings might create tiny changes in the atmosphere that may ultimately alter the path of tornado a large distance away. 

A small twist of fate can alter a lot for one person but I have always suspected the impact of  one event on the course of the entire world. Can one person, one event, one random turn, truly alter the course of history? Aren't we all just pieces in one gigantic puzzle linked together in a million tiny ways but unable to influence the picture as a whole. But then, once in a while, I encounter examples where one person does manage to alter the entire course of history - unintentionally perhaps, but irreversibly altered nevertheless. 

In many ways, the story below - laced with irony, rebellion, intrigue and love - symbolizes the Economist's law of unintended consequences. Ever since I first heard the story, I have been trying to reconcile the monumental impact of this one man, Gavrillo Princip and this one chance event on the world as it exists today. The thing that caught my attention in this story was not the action of men because they failed in their own ways. But it was the role of chance, randomness or luck that truly stood out for me. 

Gavrillo Princip (1894-1918), was a Bosnian Serb who was responsible for the assassination of Prince Archduke Ferdinand, heir to the throne of Austria-Hungary and his wife, Sophie, Duchess of Hohenburg. Princip was born in a family of serfs at a time when Serbia was in a tumultuous state of transition. 

In 1878, under the Treaty of Berlin, Austria-Hungary received the mandate to occupy and administer the Bosnia while the Ottoman empire retained official sovereignty. As part of the same treaty, Serbia was accorded the status of a sovereign state which soon transformed into a kingdom under Prince Obrenovic who ruled within the borders set by the treaty. However, this peaceful state of existence changed when as part of a military coup, the king and the queen of Serbia were violently murdered and Peter I was installed as the new king. This new dynasty was friendlier to Russia than to Austria-Hungary and over the next decade, disputes erupted as Serbia moved to reclaim its former fourteenth century empire. Serbia's military successes in these campaigns further emboldened the nationalistic elements in Serbia and the serbs in Austria-Hungary who were irked by the Austro-Hungarian rule. 

As a christian serb (serf) family living in northwestern Bosnia, the Princips (and other serbs) were often oppressed by their muslim landlords and forced to live off the little land they owned. This led to large scale discontent against the Austro-Hungrian empire. At the age of 13, Princip's brother moved him to Sarajevo and this gave him more opportunities for protest. In 1911, Princip joined the Young Bosnia, a society that wanted to separate Bosnia from Austria-Hungary and to unite it with the rising kingdom of Serbia. The following year, Princip was expelled from school for being involved in demonstrations against the Austro-Hungarian authorities. Coincidentally, after the Balkan wars in 1912-1913 the Austro-Hungarian administration in Bosnia and Herzegovina became extremely serbophobic and declared a state of emergency as the governor closed many schools and Serb societies and inflamed the historic anti-serb rhetoric. All this further fueled the young Princip and he left Sarajevo to arrive in Belgrade. He then volunteered to join Serbian Guerrilla bands fighting under the leadership of Major Vojin Tankosic, who was a member of the Black Hand - the leading terrorist organization in Serbia at the time. Three young men, including Gavrillo Princip at the age of 19, were thus trained, armed and tasked with the assassination of Prince archduke Ferdinand of Austria-Hungary by Major Tankosic. These young men were a product of their times as they sought freedom from the Austro-Hungarain empire to unite with the serbs, in hope of a better future. 




Gavrillo Princip, PC: Wikipedia 

Franz Ferdinand's life too is a charming story in itself. He was born in Austria to the younger brother of the emperor Franz Joseph, Archduke Karl Ludwig and was thus not the direct heir to the throne. However, in 1889, his cousin, Crown Prince Rudolf committed suicide and this left emperor Franz Joseph's younger brother (and Franz Ferdinand's father) next in line to the throne. When his father, Archduke Karl Ludwig, died of typhoid fever in 1896, Franz Ferdinand became the prince and heir to the throne. 

            
Archduke Franz Ferdinand, PC: Wikipedia    Duchess Sophie, PC: Wikipedia 

As a young man, Franz Ferdinand had met Countess Sophie Chotek at a ball in Prague but was forbidden to marry her as she was not a member of one of the reigning dynasties of Europe. Sophie and Prince Franz stayed in touch through letters and their relationship blossomed, away from the eyes of the court. Deeply in love, Franz Ferdinand refused to marry anyone else and after numerous appeals from him and his royal friends (Tsar Nicholas II of Russia, German emperor Wilhelm II and Pope Leo XIII all appealed his case), emperor Franz Joseph finally permitted the prince to marry Sophie. He however imposed a condition that the marriage would be morganatic and that their children would have no succession rights to the throne. Sophie was further forbidden from sharing her husband's rank, title, precedence or privileges and could normally not appear in public with him. Despite these brutal restrictions, the two married in 1900 and stayed together for the rest of the lives (and even deaths).  

In 1913, in the midst of the crisis in Serbia, Emperor Franz Joseph commanded the archduke to observe military maneuvers that were scheduled for June 1914 in Bosnia.  June was also a time of great unrest in Serbia as it commemorates the 1389 Battle of Kosovo against the Ottomans when the Sultan was assassinated by a Serb. This was a time for serbian patriotism and military observances. Although, Duchess Sophie could never share the archduke's rank and splendors as the prince; she would not let him travel alone as she feared for his safety amidst all this turmoil. In fact, if you were to believe historian AJP Taylor, love was the reason they met their deaths on this fateful day in June - "[Sophie] could never share Franz Ferdinand's] rank… could never share his splendors, could never even sit by his side on any public occasion. There was one loophole… his wife could enjoy the recognition of his rank when he was acting in a military capacity. Hence, he decided in 1914, to inspect the army in Bosnia. There at its capital Sarajevo, the Archduke and his wife could ride in an open carriage side by side…Thus, for love, did the Archduke go to his death".

On the fateful morning of June 28, 1914, The Archduke and his wife arrived in Sarajevo by train and the entire motorcade including the governor of Sarajevo began its journey as per a pre-announced program. Six armed assassins including Princip were positioned along the motorcade route with a single target in mind - Austria's heir apparent, Archduke Franz Ferdinand. 

The first two assassins along the route failed to act but the third assassin, Nedeljko Cabrinovic, who was armed with a bomb decided to take action. He threw his bomb on the motorcade but unfortunately the bomb bounced off the convertible and exploded under the next car in the motorcade. This blast caused a major furore as 16-20 people were wounded. The assassin Cabrinovic swallowed his cyanide pill and jumped into the nearby river to evade the police. Unfortunately for him though, the river was running dry and only 6 inches deep and the cyanide pill did not quite work.  He was thus taken into custody and severely beaten. A disaster seemed to have been averted as the assassination attempt appeared to have been foiled. The motorcade sped away to arrive at the town hall for the scheduled reception where the Archduke (understandably) complained about the reception accorded to him - "Mr. Mayor, I come here on a visit and I get bombs thrown at me. It is outrageous." After a few soothing words from Sophie, he finally thanked the people of Sarajevo for their ovations "as I see in them an expression of their joy at the failure of the attempt at assassination." 


After the commotion of the explosion and the rally, Franz Ferdinand and Sophie gave up their planned program and decided to visit the wounded from the bombing at the nearby hospital. The remaining assassins had all dispersed to avoid capture and it seemed that the  plot was indeed foiled. 


Now, this is where fate makes an unlikely entry and alters the course of events. 


Once the Archduke and Duchess board the motorcade, the accompanying general orders that the royal car be  taken to the Hospital through a route that avoids the city center. However, the driver of this motorcade, Leopold Lojka did not get the order and took a wrong turn into the Franz Josef street which had a cafe. Fortuitously enough, after the failed assassination attempt, Gavrillo Princip had wandered to a nearby food shop - Schiller's Delicatessen on the same street. As the universe conspired, it was at this point that the Archduke's motorcade made the wrong turn. The driver, upon being told about the changed route was trying to reverse the car when the engine stalled and the gears locked giving Princip an unexpected opportunity. Taking the chance, Princip stepped forward and fired two shots from a distance of about 5 feet. The first bullet wounded the Archduke in the jugular and the second inflicted an abdominal wound on the Duchess (who some reports say was pregnant at this time). Both victims remained seated upright but died while being driven to the Governor's residence for medical treatment. As reported by Count Harrach who was with the motorcade, Franz Ferdinand's last words were "Sophie, Sophie! Don't die! Live for our children!" followed by six or seven utterances of "It's nothing" in response to questions about his pain. 




Princip and the other assassins were meanwhile caught and imprisoned for high treason. At his sentencing, Princip stated that his second shot was aimed at Governor Potiorek than the Duchess. Princip was 19 years old at the time of the assassinations and was thus too young to receive the death penalty. In fact, he was 27 days short of his twentieth birthday which would have made him eligible for death penalty under the Habsburg law. Instead, he received the maximum sentence of twenty years in prison where he contracted tuberculosis and died on 28 April 1918. Princip had stated under cross-examination: "I am a yugoslav nationalist and I believe in unification of all South Slavs in whatever form of state and that it be free of Austria." Princip, was a young terrorist who wished for nothing but the betterment of his people. 



Assassination illustrated in the Italian newspaper Domenica del Corriere, 12 July 1914 by Achille Beltrame. 


As fate conspired, this single event - the assassination of the Archduke, triggered a chain of events that resulted in the first world war within a month. Austria-Hungary blamed Serbia and dragged Germany into the war. Russia responded and France got involved leading finally to the entry of Great Britain. The war began one month after this assassination and continued for 4 years leading to one of the bloodiest wars in world history that spanned almost the entire world in one way or another. The first world war, of course, was directly responsible for the Second world war which ultimately shaped the world as it exists today. And so, one can extrapolate that the gun shot that was intended to start a local protest by killing the Archduke actually triggered a whole lot more than that. 

Ironically, Princip was saved from the death sentence by his young age but he only lived long enough to witness the horrors of the first world war and the millions of deaths that directly resulted from his actions. I cannot be certain but I am fairly sure that given the benefit of hindsight and knowing the consequences of his actions, Princip might have chosen to not fire his gun on that fateful day. Because if you extend the chains of causation - the current middle east crisis, the Israel-Palentinian conflict, the cold war, Hiroshima and Nagasaki, Pearl Harbor, The third Reich, the treaty of Versailles, the first world war - they all occur at the other end of that one gunshot! 


I have long suspected the impact of individual agency on the course of world history because it often seems that our actions are often drowned by those of the multitudes around us, especially when it comes to changing the world. And yet, when I come across examples like this, I am forced to believe in the agency of one - for better or for worse. The agency of one, aided by the randomness and chaos that drives us is sometimes just as powerful as the flap of that butterfly's wings. 


Post-Script: 
My journey into understanding the first world war began through a podcast (Hardcore History by Dan Carlin) but it soon led me to dig deeper in books by John Keegan (The First World War) and Barbara Tuchman (The guns of August). This story has particularly captured my fascination even as I have dug-deeper and read more about it in wikipedia and other media outlets. 

Reference Sources: 
1) http://en.wikipedia.org/wiki/Archduke_Franz_Ferdinand_of_Austria
2) G. J. Meyer (2007). A world undone: The story of the great war, Bantam Dell.
3) John Keegan (2000). The First World War.
4) http://en.wikipedia.org/wiki/Assassination_of_Archduke_Franz_Ferdinand_of_Austria
5) http://en.wikipedia.org/wiki/Gavrilo_Princip
6) http://www.firstworldwar.com/bio/princip.htm
7) http://www.theguardian.com/world/2014/jun/27/gavrilo-princip-sarajevo-divided-archduke-franz-ferdinand-assassination
8) http://www.theguardian.com/world/2014/jun/27/guardian-1914-analysis-archduke-franz-ferdinand-shooting






Saturday, February 15, 2014

From being comatose to completing a marathon - all in a day! That's what happens when a python eats...

Eating a meal after a week-long starvation might be enough to make any meal seem hearty; but when a burmese python eats after starving for up to a whole year, the 'heartiness' quotient of the meal is raised to an entirely different level as the python's heart 'literally' grows by up to 40% after feeding. And, this is only one of many, quirky facts that can make the giant, scaly, unmoving, coiled hulk of a python seem interesting. 

Unlike the many snake species that actively forage and hunt for their prey, the pythons (and some others like boas, vipers and pit-vipers) employ a sit-and-wait tactic for hunting. Although this saves them the energy costs of active hunting, it also limits their feeding opportunities and imposes additional adaptations as their body has to adapt to large intervals (up to a year for a burmese python) between meals. These meals, though rare, are not small by any means - a python is capable of consuming anywhere from 80-150% of its own body weight in one meal! That's like a 100 lb. person eating anywhere from 80-150 lb. of food in one sitting. These dietary extremes have resulted in unique physiological adaptations. 

As a python lies in wait for its next meal, it enters a stage of quiescence and down-regulates both the structure and function of its gut. At the risk of damaging its own lining in the absence of food, the stomach stops producing any acid. The pancreas and the gall bladder stop their secretions and the intestinal epithelium (which normally absorbs the nutrients from a meal) atrophies. The proteins that transport nutrients across the gut lining are also shut down. In fact, in addition to the gut, other organs of a starving python such as the liver, heart, pancreas and the kidneys are also dramatically shrunk  and inactive during starvation.

But things change dramatically with that one meal. 
When a cavalier prey passes by an inert python, it is often unprepared for the viciousness of the attack and the crushing defeat that are to follow. The python lunges on its prey, impaling it with its teeth and then gradually crushing it to death by coiling around it. The muscular hulk of the python coils and crushes its thoracic cavity suffocating it to death. Once the prey is dead, then begins the slow process of swallowing it, usually head-first. The python uses its paired set of pterygoid teeth (on the palate) in alternating steps and literally walks over the prey's skull internalizing it in the process. Considering the enormous sizes of a python meal, its skull is extremely flexible and mobile with multi-hinged jaw joints and a flexible ligament between the lower jaws. The process of swallowing a large prey is further aided by the axial muscles (lining the body wall) that gently propel the food - first into the expandable esophagus and then into the waiting stomach.

Although fascinating in itself, the ingestion of the prey is only the beginning of the wondrous world of a python's alimentary canal. The prey begins to putrefy naturally soon after ingestion and generates large amounts of gas (that luckily we don't have to smell) and this further increases the girth of the python. Before you conceive of an exploding python, let me clarify that this build-up of gases is not enough to cause an explosion but it does cause severe compression and exert additional pressure on the internal organs. It also interferes with ventilation and blood flow (much more than how one might feel after two thanksgiving meals). Thus, the python needs to digest the large and intact prey before it starts decaying significantly and the only tool at its disposal is a dormant gut with no secretions. 



A Burmese python swallowing a laboratory rat which it had killed by constriction. (B) A Burmese python 24 h after consuming a rat meal greater than 50% of the snakeʼs body mass. This snake had experienced further distension of its body after feeding due to the build up of gases within the ingested dead rat.

Unlike most mammals including us, the starving python does not have baseline acid levels in its stomach. Remarkably though, within a day of the meal, the quiescent gastric lining is rapidly re-activated and it starts pumping massive amounts of H+ and Cl- ions into the stomach lumen dropping its pH from being neutral 7 at rest to extremely acidic (at pH 2) within almost 24 hours of the meal.


The post-feeding profile of gastric pH for Burmese pythons demonstrating the rapid drop in pH after feeding, the steadymaintenance of a very acidic pH during digestion, and the rise in pH upon the completion of gastric digestion when acid production ceases.

The stomach also releases an inactive protease enzyme, pepsinogen, which gets activated under the acidic conditions. The cleaved and now functional pepsinogen acts on the prey to dissolve the soft tissues and skeletal elements. By the third day, only 25% of the ingested meal (which could be as big as a kangaroo) remains within the stomach and it largely consists of the difficult-to-digest parts such as  the trunk-vertebrae, the hind limbs, tail and hair. By about 6 days, the prey is almost completely dissolved and all that remains is a mat of hair to be sent on its way out. 

                                              
                                          
Daily X-ray images of a python digesting a rat that was equal to 25% of the snakeʼs body mass. At 1 day post-feeding (DPF), the ratʼs skeleton is completely intact within the pythonʼs stomach, whereas by day 6 the ratʼs skeleton has been completely broken down and passed into the small intestine.

As the stomach acids and enzymes work, small amounts of the digested food are metered into the intestine by the opening and closing of the pyloric sphincter - a valve between the stomach and the intestine. On entering the intestine, the acidic pH of the stomach chyme is neutralized by the alkaline environment and the pH increases to 6.5 within a few centimeters. Other enzymes from the bile such as the amylases, lipase's and proteases from the pancreas also join the party as the food is broken down to its elements in the intestine. 

Once the food has been digested, the next step is the absorption of the broken-down nutrients - normally done by the tiny finger-like projections (called villi) in the intestinal wall. The python's intestine and the villi, however, are completely shriveled during the starvation period to conserve energy. However, within a few hours of the feeding (while the prey is still intact in the stomach) the dormant intestine is woken up to a flurry of activity - all in anticipation of the nutrients that will be coming in. The intestine doubles the length of the villi, up regulates the amino acid uptake rates and the activity of its enzymes.  In fact, within 24 hours of the feeding, the mass of the intestine increases by up to 70%. 
                            
Images of the small intestine of similar-sized Burmese pythons fasted and at 2 and 10 days post-feeding (DPF). By 2 DPF, the intestine has increased in diameter due primarily to hypertrophy of the epithelial cells; a response that has reversed by 10 DPF.

                       
Transmission electron micrographs illustrating the rapid post-feeding lengthening of the pythonʼs intestinal microvilli, reaching a peak in length at 3 days post-feeding. After digestion is complete (after day 6), the microvilli shorten in length and return to dormancy. Bar in images represent 1 μm or one thousandth of a millimeter.

In keeping with the increased flurry of metabolic activity, there is also an increase in the size of most, if not all, internal organs of a python - the liver, the kidneys, the heart and the pancreas. In fact, in order to keep up with the increased need for energy, nutrient transport and organ growth, the python heart grows in mass by 40% within 48 - 72 hours of a large meal. 

                              

Wet mass of the heart, pancreas, liver and kidneys plotted against time post-feeding for Burmese pythons fasted (0) and following the consumption of rodent meals equal to 25% of the snakeʼs body mass. Feeding generates respective increases in wet mass of 40%, 94%, 106% and 72% for the heart, pancreas, liver and kidneys.

The growth of an organ can be achieved by two means - by either an increase in the number of cells that make up the organ (hyperplasia) or by an increase in the size of individual cells (hypertrophy). The growth is a python heart after a meal appears to be hypertrophic or driven by increase in cell size than number. In fact, studies show that a growing python heart up-regulates a system of genes akin to what happens when we augment our cardiac output - either due to pregnancy or with exercise. These events are rather different from the pathological program that is activated by hypertension, heart attacks etc. The quirky physiology of a python heart thus becomes an interesting subject to study even from the perspective of human health and disease. 
                  
                                      

The increase in the size of a python heart post-feeding is characterized by cellular hypertrophy. Compare the heart size of a starving python and that at 3 days post-feeding (DPF) Scale bar, 2 mm.

Two days after feeding, after absorption of the nutrients, the unabsorbed material (largely hair) begins to enter the large intestine. With each continuing day of digestion, the cecum and the large intestine fill with unabsorbed waste material, that is ultimately expunged as a bolus along with urate. Within a week after the meal, the last of the prey exits the stomach and the small intestine and these organs  begin the shut-down process anew. Within ten days, the stomach pH rises, the enzyme activity drops down and the intestinal mass declines. The other organs too begin to shrink to their former state.

The question that remains though is what regulates this system level change in the organ size and function. 

Studies suggest the involvement of two pathways - hormonal (systemic effect of hormones) and luminal (arising from the gut lumen itself) in mediating these rapid changes in organ size and function. The levels of many hormones that regulate gut function such as neurotensin, glucagon, insulin and cholecystokinin increase sharply after a meal. Experiments where small segments of the gut were surgically isolated from the rest of the gut but kept connected to the circulation were also able to increase their function upon a meal suggesting that a link to the blood circulation was sufficient to drive these systemwide changes. The gut lumen itself was also seen to play a role in triggering the initial responses as direct injection of nutrients (amino acids and proteins) into the python's small intestine was able to reawaken the intestine from dormancy. Interestingly though, injecting saline, glucose or lipids did not have the same effect on the python's intestine. 

The factors underlying the increased heart size are however much less clear. In order to study the possible involvement of systemic factors on cardiac output, scientists injected blood plasma from fed and starving pythons into mice. Much to their surprise, components in the plasma of a fed python had similar effects on the heart of a starving python and of a mouse and caused significant increase in their respective heart sizes. 
                               

Fatty acids in the blood of a fed python induced cardiac growth in a starving python. Infusing fasted pythons with fed plasma or a mixture of the three individual fatty acids resulted in increased heart mass (heart weight/body weight) thus mimicking the effect of a meal (3 DPF). Bovine serum albumin (BSA) was used to solubilize FAs and is therefore used as control here to show that it alone did not have the safe effect as the fatty acids.

                                                              
Seven-day infusion of the same three fatty acids in mice also resulted in increased heart growth (as measured by the size of the ventricle).

Interestingly, the fed-python-plasma continued to function in a starving python even after treatment with heat and a protease - suggesting that the active ingredient was likely not a protein. Scientists then suspected the involvement of lipids (since they are resistant to heat and proteases) since feeding led to sharp increases in the levels of lipids and fatty acids in the python blood (see the turbid opaqueness of python plasma after feeding). Careful studies identified three commonly known lipids - myristic, palimitic and palmitoleic acid to be responsible. Injecting these three lipids in the precise ratio to a starving python over a 7 day period resulted in significant growth in the heart - comparable to what would have been triggered by a good meal. Strangely however, the effect of these lipids was limited to the heart as the other organs - liver or muscle showed no change. Supplementing these lipids individually also had no effect. 
                           

Levels of non-esterified fatty acid (NEFA) and triacylglyceride (TAG) concentrations in the blood plasma are significantly increased after feeding (see the milky opaqueness of the plasma at day 1 - 2 after feeding).


Careful observation also shows that the levels of the lipids shoot up in the plasma even before the digestion has begun in earnest (see peaks at days 1 and 2 and compare with the X-rays of the prey from earlier). This suggests that the lipids are not coming from the digestion of the prey but are indeed a signal produced by the python to synchronize its various organ systems and to prepare them for the upcoming phase of hyperactivity. 

Also of interest is the fact that despite such high levels of lipids in its blood (to the extent that the plasma appears cloudy), the python heart shows no lipid deposition. This again would be rather useful to many of us who worry about inching closer to a heart attack with every blob of butter or every additional french fry. If only, our hearts would be more like a python's and not deposit the fat around its vessels - we could eat everything that we wanted and yet avoid the consequences (or at least some of them).  

Despite the creepy-crawlies that I had previously associated with a python, now, it is a fascinating creature with some extremely cool biology. Imagine someone emerging from a near comatose state only to successfully complete a marathon! That is the equivalent of what a python does every time it eats a meal. 

References:

1) Cecilia A. Riquelme et al. Cardiac Growth Fatty Acids Identified in the Burmese Python Promote Beneficial cardiac growth in Science 334, 528 (2011)

2) Digestive physiology of the Burmese python: broad regulation of integrated performance
by Stephen M. Secor The Journal of Experimental Biology 211, 3767-3774 in 2008



Thursday, December 26, 2013

The glass menagerie

At the end of two flights of stairs, in the middle of the bustling Harvard campus, lies a time capsule that few are aware of. On a grey morning in June, I walked up the dark, metal stairway for my first encounter with this well-kept secret - 'The glass menagerie'. 




I stood surrounded by victorian cherry wood vitrines proudly displaying nearly 4,300 three-dimensional botanical specimens, representing nearly 840 species of plants from a 170 different families. In addition to their evergreen and brilliantly natural hues, they are also pristinely delicate, fragile and over a century old. They are not a modern technological marvel; they are in fact a relic of history. From a time when scientific study of the natural world was plagued by the limitations of time and resources, and when practical matters of distance, transport, storage and preservation could not be trivialized. 

These specimens are the part of a glass menagerie that is more than a century old. Their's is a tale of wonder that is shaped by the primal human obsession to collect and create that and the narrative begins in the Renaissance Europe. 

In the sixteenth century, Europe woke up after the dark ages to the rest of the world in a period of renaissance. Status-conscious royals, nobles, physicians and apothecaries - anyone who could afford to - began assembling eclectic objects in a single room. 'Wunderkammern' or cabinets of curiosity, as they were called, are the ancestors of our modern day museums. They broadly expressed the beautiful, the monstrous, and the exotic: preserved flora and fauna, scientific instruments, objects of art and genetic mutations. They began as odes to idiosyncrasy driven by an obsession to collect but soon transformed into precursors of a  scientific quest that goes on till today. 

One of the earliest steps in this transformation from the 'weird' to the wonderful was the development of the universal classification system in 1753 by Carolus Linneaus (1707-1778), a swedish botanist. Linneaus believed that, "The first step in wisdom is to know the things themselves" and thus devised a simple, beautiful and instructive way to classify all living things using two word names in latin - first identifying the genus, and the second, the species. His work was carried forward by biologists such as Georges Cuvier (1769-1832) and Jean-Baptiste-Pierre-Antoine de Monet de Lamarck (1744-1829). Their research led to a flood of publications. Some of them, like Pierre Joseph Redoute's Les Liliacees, Les Roses and James Audobon's Birds of America were not only scientific treatises but also masterpieces of draftsmanship and printing. 

Linnaeus and the Enlightenment also paved way for proper scientific collecting as naturalists began to prepare their specimens with greater care. But early preservation techniques were crude and did harm than good as insects were pickled in spirits, snakes were crammed with straw, shells were boiled and shipped in sawdust. Naturalists were thus reduced to studying animals from illustrations and text books that were painstakingly made to try and recapitulate nature's three dimensional wonders on flat two dimensional plates that gave no indication of size or scale and were open to misinterpretation. 

Around the same time, England was also caught in the throes of another craze, as Philip Henry Gosse (1810-1888), a self taught British naturalist popularized the notion of keeping sea creatures in oxygenated saltwater aquariums. His Actinologica Brittanica: A history of British sea-anemones and corals was a compendium of his illustrations by keeping animals in such aquariums. The British craze for aquariums began from the idea of 'Wunderkammern' but was fueled by the availability of inexpensive glass plates and the discovery that sea weed could be used to oxygenate water in the aquariums. At this time, the generally successful method of transporting live specimens of sea anemones and mollusks over long distances consisted of wrapping them in wet seaweed, placing them in glass jars and packing the jars in baskets. With these and other improvements, the first large public aquarium was established by the Zoological Society of London in 1853. 

Around the same time (between 1872 and 1876), in keeping with the spirit of exploration, the British Admirality and the Royal Society also initiated the Challenger expedition - a massive feat of oceanographic exploration where the HMS Challenger, a massive 2300 ton warship, covered nearly 69,000 nautical miles charting the world's oceans as oceanographers mapped the seabed and ocean currents and biologists collected thousands of species of marine life. The findings of this expedition were studied by prominent researchers of the time like Ernst Haeckel (1834-1919) and were published as a compendium in 50 volumes. 

And thus the quest that began as a private luxury in the homes of the affluent, soon transformed into a public fascination as museums began opening their doors to the public in the late 18th and early 19th centuries. Subsequent to the Louvre in 1793 and the Prado in Madrid in 1809, public museums began mushrooming all over Europe and America. In this climate of scientific wonder and renaissance began the story of the glass menagerie and its two creators - the father-son duo of the Blaschkas. 



























The story of the Blaschkas begins in the small town of Bohmisch Aicha, in the now Czech Republic, where Leopold Blaschka, the father, continued his family tradition of flame working. Despite a strong interest in natural history and art, Leopold entered the family business of making costume jewelry and other fancy goods with metal and flame worked glass. His quaint world was however shattered by two devastating losses when his first wife died of Cholera in 1850, followed by his father in a couple of years. Devastated, the grieving young Leopold took time off to visit the United States. On his maiden voyage though the ship was becalmed for two weeks near the Azores and young Leopold passed his time collecting and illustrating jelly fish and other marine invertebrates. Their glasslike transparency fascinated this flame worker and this sense of wonder transformed his life from there-on. 

Leopold describes his sense of awe and wonder (in this translation provided by Henri Reiling) -
"It is a beautiful night in May. Hopeful, we look out over the darkness of the sea, which is as smooth as a mirror; there emerges all around in various places a flash like bundle of light beams, as if it is surrounded by thousands of sparks, that forms true bundles of fire and of other bright lighting spots, and the seemingly mirrored stars."

After the death of his wife, Leopold had sought consolation in collecting, studying and painting plants and upon his return to Europe after his wondrous encounters with the glass-like creatures, he began making glass models of plants for his own amusement.  These models came to the attention of Prince Camille de Rohan, an aristocratic horticulturist who has established a world-famous garden on one of his estates at the Sychrov castle near Aicha. Between 1860 and 1862, the prince exhibited nearly 100 models of orchids and other exotic plants - all in glass. 


Leopold's journey began with making costume jewelry, chandeliers and other fancy goods but gradually expanded into jewelry decorated with flame worked flowers, flame worked glass eyes and laboratory equipment. However, his work with the Prince opened new avenues and led him onto a new vocation - the art of making scientific models.  The Blaschkas' models varied greatly in complexity and in their method of construction. Component parts were formed from both clear and colored glass using a combination of lampworking and glassblowing. The parts were then either fused together or assembled with adhesives, probably hide glue. Where necessary, other materials were used in the construction: fine copper wires were added to reinforce delicate tentacles and gills and painted paper was cleverly incorporated to represent internal structures. Surfaces were painted with colors mixed with gum or glue.

Impressed with Leopold's craftsmanship, the prince introduced Leopold to Prof. Ludwig Reichenbach, director of the botanical garden and the natural history museum in Dresden and this resulted in regular exhibitions of Leopold's glass models in the city's botanical garden and in a museum in Liege, Belgium. Prof. Reichenbach then commissioned Leopold to make models of sea anemones and other marine invertebrates to be displayed at the natural history museum and this further attracted the attention of other museum directors. 

By the age of 40, Leopold was a successful model maker. His skilled flame working, apprenticeship with a jeweler and years of running a family business had amply trained him for working at a very small scale. Leopold used several design sources beginning with the printed page from the works of PH Gosse (who popularized the aquarium). Gosse's illustrations provided Leopold with images of sea anemones and also suggested that the models could be displayed on natural, rock-like surfaces. Gosse's plates however gave no sense of scale or dimension which an accurate three dimensional model needs and this set Leopold on the lookout for other sources. Around this time, Leopold was also joined by his son Rudolf, who brought fresh zeal and enterprise to this venture. 

Over the years, the Blaschka duo worked from books and other publications from all over Europe. True to the times, they also relied on animals preserved in glass jars of alcohol but these were a mixed blessing as the specimens lost their coloring and having no backbones to support them collapsed into shapeless masses at the bottom of the jars. Rudolf was an enthusiastic assistant to his father and soon injected new ideas into the business. The Blaschkas began to maintain living specimens in seawater aquariums of the kind promoted by Gosse and could successfully maintain anemones for 'years'. They acquired live specimens from Naples, from Chioggia and Trieste in the upper Adriatic, from Weymouth on the english channel and from suppliers on the coasts of the North and Baltic seas. Leopold further expanded these sources by venturing on expeditions to document new species. In 1879 for example, Leopold went on an field trip to the upper adriatic which afforded him an opportunity to observe a greater number and variety of invertebrates. Later, he made more ambitious field trips to the United states and the Carribbean in 1892 and 1895. 





The Blaschka's glass models were well timed with the aquarium craze that swept through England and they made it possible to stock waterless aquariums with sea anemones and other invertebrates. These required little or no maintenance, and unlike true aquariums, no restocking in event of death. The glass models also retained their shape and color. The Blashckas' archive of drawings and sketches from textbooks further ensured that these models were accurate. The glass anemones were soon joined by corals, jellyfish, mollusks and other species. 

The Blaschkas' skill in producing minutely detailed replicas was further supported by the socio-political climes of the era. Beginning with the French revolution, traditional values were challenged and in many cases transformed in Europe. The development of science and the expansion of public education provided greater opportunities for their models. The Blashckas' succeeded because their models solved a problem that confronted all directors of Natural history museums. While the vertebrates could be displayed relatively easily by stuffing and mounting, the taxidermists could not work their magic on the invertebrates (jellyfish, squids and so forth). As opposed to the limited scope of the existing method of bottles of alcohol, the glass sculptures provided museum curators with displays of permanent form and feature. 









The proliferation of museums and the resultant emergence of suppliers to these museums, expanded the Blaschkas' presence. They soon had a presence in India, New Zealand, Tokyo, Austria, Ireland, Scotland, Germany, Belgium, Netherlands, Switzerland, Australia, France and multiple places in the United States. The father-son duo managed to maintain a prodigious output through the years as they made hundreds and thousands of models. 

Harvard university, my haunt for the day, began forming its teaching collections in the 1850s under the leadership of zoologist, Louis Agassiz (1807-1873). At the founding of the museum of comparative zoology in 1858, Agassiz made an impassioned speech about the lack of a teaching collection in the United States which necessitated the students traveling to Europe. He vowed to remedy the situation and it was Agassiz who acquired the zoological museum's first 350 Blaschka models. 

In 1886, George Lincoln Goodale, a professor of Botany at Harvard University, traveled to Germany to persuade Leopold to abandon his successful career making models of invertebrates and to focus on plants instead. As the director of the Harvard Botanic Garden, Prof Goodale desired to represent the full glory of the plant kingdom in the natural history museums that were being developed at Harvard. The display would also help supplement the botanical courses as many plant beds and greenhouses were subject to the inhospitable New England winters, further limiting the available options. Prof. Goodale was on the lookout for something aesthetically pleasing and scientifically accurate, since presenting plants as attractive displays was tougher. Traditional botanical teaching aids included models fashioned from wax-covered silk or Papier mache in addition to fresh fruits and flowers. Although dried, pressed herbarium specimens were also frequently used to supplement these options, these had limited appeal and utility. Impressed by the glass zoological models at the museum, Prof. Goodale desired something similar for his plants to point out their morphological features during lectures. He thus traveled to Dresden, Germany where the Blaschkas lived and ultimately persuaded them to accept a small commission for a few plant models. This in some sense brought Leopold's journey a full circle and the result was a unique collection of botanical models, that I stand witness to today. These became known as the glass flowers of Harvard and while the invertebrate models are spectacular, the botanical models are simply breathtaking for their sheer diversity and accuracy. 

With Prof Goodale's advocacy and generous financial support from Mary Lee Ware, the initial small commission to the Blaschkas was extended into a ten year contract and the resulting glass models were titled the Ware collection of Blaschka glass models of plants. This collection represents the diversity of flora, with an emphasis on economically important plants used in everything, from food to medicines. The models were largely based on plants that the Blaschkas cultivated on their property from various sources. By the mid-1890s, the Blaschkas had made models of several hundred species to represent the major plant families and had begun to exhaust their european sources. 

At this point, Rudolf was anxious to visit America to study some of the plants of interest in their native environment and to make fresh studies. Armed with color pencils and drawing paper, Rudolf began his voyage to America early in 1892. Rudolf began a diligent study of the flowering plants in the Harvard botanic garden and also completed an expedition to Jamaica. He also traveled the vast North American continent, extensively sampling the terrain and the interesting flora. In these voyages he made extensive drawings for himself and his father. Unlike the invertebrate models, Rudolf drew the plans in their natural settings and also dissected them in order to highlight their various parts, either life size or magnified. He made detailed illustrations to provide reference information on color, and dimensionality that would be lost from the herbarium specimens as the plants were pressed and dried. He also included top, front, back and side views of the flowers and their internal structures such as stamens and pistils. He drew cutaway views to record the size ratios and the placement of each part of the plant. His notes also provide critical information about the plants such as the number denoting the color (to try and correlate with the pigments used by his father to paint the parts of the colored glass models). He also gave descriptors for textures, sheen, opacity etc in addition to designations for twists, furrows, creases, wrinkles and spots.  

Although the surviving archive of specimen, drawings and labels is incomplete, Rudolf probably documented upwards of 350 species, and with Leopold created more than 250 sets of models based on this 1892 season alone. The work proved so valuable that he returned for a second season in three years - this time as a more experienced traveler. It was during this second season of exploration that Leopold suffered a stroke and died leaving Rudolf with the arduous task of finishing the models that Leopold had begun. 















Despite the lack of any formal training, Rudolf was thorough and meticulous as any scientist in the day as he recorded the unique or relevant features of a genus or a species in addition to the physical characteristics. From 1896 to 1936, when the last shipments of the models were received, he added more than 200 species, including several series on grasses, insect pollination, progression of fruit blight etc. Rudolf Blaschka continued his work through his final years and despite his failing health, his devotion and fascination to these models remained unaffected. Unfinished models remained on his desk when he died on May 1, 1939. 

Rudolf writes in a letter to an American colleagues, Walter Deane in 1899:

"I think I belong to that same order of men as you, to the true lovers of nature. On every walk I take, there must be something to study of nature, it maybe a plant or insect or bird or whatever. I think a man can never finish these studies and is never too old to learn from nature..... What I saw and learned from nature on those trips in America gives me very sweet hours of remembering for all life." 

This glass menagerie has its origins in Europe but continues to fascinate museum-goers all over the world today. It is a legacy of Leopold and Rudolf Blaschka and while it does offer insights into the history of science and the perspectives on model-making itself, it also reveals a delicate and often invisible link between artists and scientists - like the glass it is made of.